1\/IBL/WH01 THE NAUTILUS Volume 109 1995 AUTHOR INDEX Anseeuw, P 117 CoovERT, G. A 43 CoovERT, H. K 43 GlTTENBERGER, E 38 GOFAS, S 14 Goto, Y 117 Groves. L. T 113 Kreiser, BR HI Martins, A M. F 1 MiTTON, J. B Ill Povel, G. D E 38 Shen, J 41 Squires, R L 113 TippETT, D. L 127 Vermeij, G. J 120 NEW TAXA PROPOSED IN VOLUME 109 (1995) GASTROPODA Pleurotomariidae Perotrochus metivieri .\nseeuw and Goto, 1995, new species 117 Barleeidae Barleeia aemilii Gofas, 1995, new species 19 Pseudodiala aequinoctialis Gofas, 1995, new species 26 Barleeia chefiae Gofas, 1995, new species 19 Barleeia cinguloides Gofas, 1995, new species 22 Pseudodiala corollaria Gofas, 1995, new species 26 Lirobarleeia elata Gofas, 1995, new species 28 Pseudodiala niso Gofas, 1995, new species 24 Barleeia pervulgata Gofas, 1995, new species 20 Barleeia picta Gofas, 1995, new species 24 Lirobarleeia pupoides Gofas, 1995, new species 29 Lirobarleeia sublaevis Gofas, 1995, new species 28 Barleeia taeniolata Gofas, 1995, new species 20 Barleeia tomensis Gofas, 1995, new species 20 Barleeia verdensis Gofas, 1995, new species 17 Cypraeidae Proadusta goedertorum Groves and Squires, 1995, new species 114 Cystiscidae Granulininae Coovert and Coovert, 1995, new subfamily 73 Persiculinae Coovert and Coovert, 1995, new subfamily 70 Plesiocystiscinae Coovert and Coovert, 1995, new subfamily 66 Plesiocystiscus Coovert and Coovert, 1995, new genus 66 Marginellidae Austroginellini Coovert and Coovert, 1995. new tribe 73 Serrataginella Coovert and Coovert, 1995, new genus 82 Prunini Coovert and Coovert, 1995, new tribe 89 Turridae Drillia fClathrodrillia) dautzenbergi Tippett, 1995, new name 129 Clalhurella eversoni Tippett, 1995, new species 129 Feniinorea kathijae Tippett, 1995, new species 132 Fenimorea petiti Tippett, 1995, new species 132 Drillia (Clathrodrillia) petuchi Tippett, 1995, new species 133 Sediliopsis riosi Tippett, 1995, new species 133 Drillia (Drillia) wolfei Tippett, 1995, new species 127 Ellobiidae Allochroa nana Martins, 1995, new species 8 Allochroa tenuis Martins. 1995, new species _ 8 THE NAUTILUS Volninc 109, Niimhcr 1 August 31, 1995 ISS\n()2.S-1344 A (juartcrhj devoted la nudacologij. EDITOR-IN-CHIEF Dr. M. G. HarascwTcli Division of Molliisks National Museum of Natural History Smithsonian Institution Washington, DC 20560 ASSOCIATE EDITOR Dr. R.Tucker Ahhott American Malacologists, Inc. RO. Box 2255 Melbourne, FL 32902 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Arthur E. Bogan Freshwater MoHuscan Research 36 Venus Wdv Sevvell, NJ 080S0 Dr. Robert T.Dillon. Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. Wilhani K.. I'hucrson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Mr. Richard I. jolmson Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Aurele La Rocque Department of Geology The Ohio State University Cohiinbus. OH 43210 Dr. James H. McLean Department of Malacology Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department ol Malacology Delaware Museum of Natural Histon RO. Box .3937 Wilmington, DE 19807 Dr. Donald R. .Moore Division of Marine Geologv and Geophysics Rosenstiel School of Marine and Atmospheric Science University oi Miami 4600 Rickcnbackcr f ^ansrwax Miami. FL 33149 Dr. Gustav Paulav Marine Laboratoiy University of Guam Mangilao'. Guam 96923 Mr. Richard E. Petit RO. Box .30 North Myrtle Beach, SC 29582 Dr. EdwardJ.Petuch Department of Geologj' Florida Atlantic University Boca Raton, FL. 33431 Dr. David H. Stansberv Museum of Zoology The Ohio State University Columbus, OH 4.3210 Dr. Ruth D. Turner Department of .Mollusks Museutu of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Geerat J. 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MD 20907-7279 THEe^NAUTILUS CONTENTS Volume 109, Number 1 August SI, 1995 ISSN 0028-1344 Antonio M. de Frias Martins Systematic Revision of Atlochroa Ancey, 1887 (Pulmonata: Ellobiidae) with an Account of the Anatomy of Allochroa layardi (H. & A. Adams, 1855) and the Description of Two New Species 1 A Remarkable Species Richness of the Barleeidae (Gastropoda: Rissoacea) in the Eastern Atlantic 14 Shell Growth and Decollation in Terrestrial Gastropods 38 Cannibalism in the Terrestrial Slug Deroceras lacve 41 Serge Cofas Edmund Gittenberger G. David E. Povel Jianming Shen THE NAUTILUS 109(1):1-13, 1995 Page 1 Systematic Revision of Allochroa Ancey, 1887 (Pulmonata: Ellobiidae), with an Account of the Anatomy of Allochroa layardi (H. & A. Adams, 1855) and the Description of Two New Species Antonio M. de Frias Martins Dt'pailaiiU'iito tie Biologia Universidade dos A(,ores P-9502 Ponta Delgada codex Sao Miguel, A(;ores, Portugal ABSTRACT Of the nominal species of Allochroa Ancey, 1887 described to date, only two are recognized: A. hronnii (Pliilippi, 1846), en- demic to the Hawaiian Islands, and A. layardi (H & A Adams, 1855), distributed throughout the Indo-Pacific. Examination of the I'SNM collection revealed two additional species, A. nana Martins, n. sp. and A. tenuis Martins, n sp,, herein described. The anatomy of A. laijardi is described and serves as the basis for determining the relationships of this genus within the El- lobiidae. Allochroa has a monaulic, entirely glandular pallial gonoduct and a wide visceral nerve ring with a long right parietovisceral connective, confirming placement in the Py- thiinae Its close relationship to the Mediterranean and Eastern North .\tlantic genus Ovatella Bivona, 1832 is indicated by similar apertural dentition, protoconch morphology of A. ten- uis, and b\ the internal structure of the penial complex, where the penis appears as the development of one of the pilasters. Key Words: Ellobiidae, Pythiinae, Allochroa, systematics, anatomy. INTRODUCTION The Ellobiidae are a group of primitive, mostly halo- philic pulmonale mollusks, commonly found near and above the hightide limit of mangroves, salt marshes, and boulder and cobble beaches. Morphologically very di- verse and ranging in length from barely 1 mm (Leu- conopsis ) to about 100 mm (Ellobium). they nevertheless share basic shell characters and body features that allow them to be considered within a single family. Pfeiffer (1853) was the first to attempt a subfamilial organization, based on shell characters alone. He assigned those species with a thick, reflected outer lip to the Auriculea [=EI- lobiinae], while he included the species with sharp outer lip in the Melampea [=Melampodinae]. Utilizing radular and also some anatomical characters, Odhner (1925) rec- ognized a more complex subfamilial arrangement and redistributed the ellobiid genera among four additional subfamilies: Carychiinae, Pedipedinae, Pythiinae and Cassidulinae. Odhner s outline was supported on con- chological features by Zilch (1959). The first detailed use of anatomical characters, in a review that included spe- cies belonging to 11 genera, was by Morton (1955). His study supported Odhner's classification, but the Cassi- dulinae were included in the Pythiinae, on the basis of similarities of the reproductive system. In a study of species belonging to 17 of the 23 genera currently rec- ognized in the Ellobiidae, Martins (in press a) clarified Morton's (1955) findings by redefining the subfamilial morphological patterns of the reproductive and nervous systems. Preserved specimens of Allochroa layardi (H. & A. Adams, 1855) are used here as the basis for detailed anatomical investigations of the family extending the preliminary findings reported in Martins (in press b). Insight on the anatomy of A. layardi provided infor- mation needed to define the phylogenetic relationships of Allochroa Ancey, 1887 within the Pythiinae. The pres- ent work reviews the taxonomy of the species of Alloch- roa and describes the anatomy of A. layardi. Examina- tion of the Mollusk collection at the National Museum of Natural History, Smithsonian Institution, consisting of only dry material, revealed the existence of two new species, herein described. MATERIALS AND METHODS This taxonomic revision is based primarily on dry spec- imens of Allochroa in the USNM collection. Four spec- imens of A. layardi, collected on Henderson Island in 1991, were provided for anatomical work. The fully re- tracted animals were preserved in alcohol and, except for the eyes and the tip of the tentacles, appear to have lost all other pigmentation. Two specimens were dis- sected under a binocular microscope and drawn using a camera lucida. Shells with protoconchs were mounted using double- Page 2 THE NAUTILUS, Vol. 109, No. 1 coated tape. Radulae were stretched across a cover slip after cleaning with KOH solution. Both structures were mounted on stubs and examined and photographed using a Cambridge Stereoscan 250 Mk2 Scanning Electron Mi- croscope. Acronyms for specimen repositories are as follows: ANSP: The Academy of Natural Sciences, Philadel- phia BMNH: The Natural History Museum, London MCZ: Museum of Comparative Zoology, Harvard LIniversity, Cambridge MNHN: Museum National d'Histoire Naturelle de Paris USNM: National Museum of Natural History, Smith- sonian Institution, Washington, D.C. KEY TO THE SPECIES OF ALLOCHROA ANCEY, 1887: 1. Shell thick 2 Shell thin .'3 2. Aperture oval elongated (base narrow) A. bronnii Aperture auriculate (base expanded) A. layardi 3. Shell length at least 4 mm 4 Shell length less than 4 mm 5 4. Protoconch width less than .5 mm A. layardi Protoconch width at least .6 mm 5 5. Protoconch without umbilicus and with apertural lip not reflected A. nana Protoconch with umbilicus and with apertural lip slightly reflected A. tenuis SYSTEMATICS Class GASTROPODA Cuvier, 1797 Subclass PULMONATA Cuvier, 1817 Order Archaeopulmonata Morton, 1955 Family Ellobiidae H. & A. Adams in Pfeiffer, 1854 Subfamily Pythiinae Odhner, 1925 Genus Allochroa Ancey, 1887 Allochroa Aiiiey, 1887:288 Type species by original designa- tion: Auricula bronnii Pliilippi, 18-1(1 Description: Shell lengtli to 12 mm, oval-conic, fragile to .somewhat solid, sometimes hirsute. Umbilicus absent Spire with up to 7.4 flat to slightly convex whorls, sculp- tured with more or le.ss marked spiral lines; body whorl 55 to 80% of shell Icngtli, usually with three chestnut spiral bands over a whitish to light-brown background, the band just above the aperture consistently strongest, the one near the suture sometimes faint, the one near the columella largest, sometimes coalescing w ith the me- dian band. Aperture about 50 to 60% of shell length, oval-elongated to .semi-circular, angled posteriorly, rounded anteriorly; inner lip slightly reflected over the columella, with three more or less etiuidistant teeth: one oblique columellar tooth, a more or less horizontal an- terior parietal tooth, and a sometimes strongest, oblique posterior parietal tooth; outer lip sharp, usually with one prominent fold entering the aperture at about the level of the parietal teeth, forming a somewhat marked anal canal. Protoconch bulbous, smooth, with up to 1.3 whorls visible. Radula: Central tooth narrow, base deeply emargin- ate, with two median flaring arms, unicuspid; lateral teeth with elongated, sinuous base with two projecting asymmetric arms, inner arm lower, crown large and long, subquadrangular; marginal teeth with sharp endocone gradually becoming as long as mesocone as row pro- gresses outward, developing a sometimes conspicuous basal ectocone. Anatomy: Reproductive system monaulic, pallial gon- oducts entirely glandular, pallial gland present. Central nervous system with wide visceral ring; right parieto- visceral connective as long as left. Remarks: The name Allochroa was proposed by Ancey (1887) to replace the supposedly preoccupied "Laimo- donta des auteurs [H. & A. Adams, 1853] (non Nuttall)" (type species Auricula Inonnii Philippi, 1846). "Lai- modonta Nuttall' , wrongly taken as an emendation of Laemodonta Philippi, 1846, was restricted to a different assemblage (type species Auricu/as/ria^a Philippi, 1846), following Nevill (1878). A more detailed account of the synonymy of Laemodonta and Laimodonta is presented elsewhere (Martins, in press a). Ancey (1887) also made clear that Marinula King, 1832, very similar to Pedipes Scopoli, 1777, was not an eligible group for his new assemblage, and that Monica H. & A. Adams, 1855[=Oiw^e'//a Bivona, 1832], although decidedly conchologically closely related, is restricted to Mediterranean and West Atlantic species. Allochroa is conchologically characterized by the pres- ence of an outer lip internal lamella which, in conjunc- tion with the anterior parietal tooth, divides the aperture into two roughly equal halves: a posterior portion, where the shell is thinner; an anterior, thickened half, some- times with an additional internal fold on the outer lip. The inner lip ol the aperture is typically pythiine, i.e., roughly equidistantly tridentate, with one small colu- mellar tooth and two parietal teeth. The lamellar, outer lip tooth readily separates this genus from the closely related Laemodonta, which usually has two round tu- bercles on the outer lip, and also from Ovatella, where the outer lip lamellae are much shorter. The single col- umellar tooth sets Allochroa apart from Pedipes which, as in P. mirahilis (Miihlfeld, 1816) and P. pedipes (Bru- guiere, 1789), has a strong, crest-like outer lip fold pen- etrating inside the aperture. In Pedipes, the single pa- rietal tooth is very strongly pronoimced. The six s])ecies assigned by authors to Allochroa are here synonymized into two: the Hawaiian Allochroa Inonnii (Philippi, 1846) and the Indo-Pacific Allochroa layardi (H. & A. Adams, 1855). Two additional new species are described herein. Kuroda and Habe (1952) listed Pedipes ajfinis F"er- ussac, 1821, under A//of/jroa, owing prol)ably to the pres- ence of a strong lamella inside the outer lip. Ferussac A. M. de F"rias Martins, 1995 Page 3 (1821), however, had mentioned a bidentate cohimella as characteristic of Pedipes, and Ancey (1887) also used this feature of P. affinis to separate it from Laernodonta , where Jickeli (1874) had erroneously placed it. Following the above reasoning, I concur with Ancey (1887) in con- sidering Ferussac's species a Pedipes. Similarly, the New Caledonian Pedipes forestieri Sou- verbie and Montrouzier, 1864, has been erroneously as- signed to Allochroa (Franc, 1954; Smith, 1992). In the original description Souverbie and Montrouzier (1864a) mentioned the presence of two columellar teeth, stating that their observations were based on a single, probably juvenile specimen. In a later paper (1864b) the same authors, reacting to the epistolary comment of L. Pfeiffer on that subject "Spec, tiov., sed Marinida". changed the ta.xonomic combination to "Marinula (an Pedipes?)", cautiously justifying, however, their original generic choice by pointing out features such as " 1 ) shell positively striated transversely; 2) two positively columellar folds; 3) external border of the columellar lip contouring be- hind the upper columellar fold as in Pedipes mirabilis, Miihlf., which we possess." The accompanying figure (Souverbie & Montrouzier, 1864a: pi. X, fig. 1) appro- priately illustrated their reasoning. Franc (1954) claimed to have found an adult specimen of Souverbie and Mon- trouzier s species in the Jousseaume collection at the MNHN, and assigned it to Allochroa. The redescription of Pedipes forestieri provided by Franc (1954), mentions the presence of a spiral fold inside the outer lip char- acteristic of Allochroa, but corroborates the presence of "two columellar folds closer to each other than to the parietal", a feature diagnostic of some pedipedine gen- era, namely Pedipes and Marinula. Taking into account Souverbie and Montrouzier's (1864a, b) insistence on the pedipedine characteristics of the apertural dentition, a feature confirmed by Franc (1954), and in my inability to examine the type material, I consider the assignment of Pedipes forestieri to Allochroa to be unwarranted. The similarity of the basic patterns of shell morphology and color throughout the genus creates some difficulties in separating species. Larger specimens of the highly variable, widely distributed A. layardi overlap in shape and color with smaller specimens of the Hawaiian A. hronnii (compare Figs. 1 and 5). At the other extreme, thin, less distinctly banded forms of A. layardi could be confused with lighter specimens of A. tenuis, n. sp. (com- pare Figs. 10 and 19) However, species can be readily separated on the basis of other conchological characters such as the ratio number of whorls/shell length or using protoconch features. Allochroa bronnii (Philippi, 1846) Figures 1-2, 21, 31-32 Auricula Irronnii Philippi, 1846;98 [Hawaiian Islands; location of type unknown]. Auricula sandwichiensis Souleyet, 1852: 524, pi. 29, figs. 29-32 [Hawaiian Islands; type not found, supposedly at BMNH (Gray, 1855)]. Ophicardelus (Lai- modonta) sandwichensis (Eydoux & Souleyet). H. & A. Adams, 1855a: 34 [misspelling of santftuic/iiensis]. Melam- pus bronni (Philippi). Pfeiffer 1854:147. Ellobium hrownii (Philippi) H & A Adams, 1855b:237 [misspelling of fcron- nii] Laimodunta sandwichensis (Eydoux & Souleyet). H. & A Adams. 1855b:246, pi. 83, figs 1, la [misspelling of sandwichiensis]. Mclampus sandwichiensis (Souleyet). Pfeiffer, 1856:50. Ophicardelus sandwichensis (Eydoux). J. E. Gray in M. E. Gray, 1859:(3), pi. 304, fig. 2; (4):1I8. Laernodonta Inonni (Philippi). Martens & Langkavel, 1871: 57. Marinula {Laernodonta) bronni (Philippi). Nevill, 1878: 220. Allochroa Inonni (Philippi). Ancey, 1887:288. Tralia {Laernodonta) Inonni (Philippi). Crosse, 1894:321. Me- lampus (Laimodonta) Ifronnii (Philippi). Kobelt, 1898: 200, pi. 23, figs. 1, 2. Laimodonta bronni (PhiMppi). Dautz- enberg & Fischer, 1905: 107 Description: Shell (Figs. 1-2, 21) length to 12 mm, ovate- conical, somewhat solid; violet-brown to light-brown, banded. Spire moderately high with up to 7.4 flattened whorls, with marked spiral striae crossed by irregular growth lines that give the surface a matte appearance; one light-brown subsutural band, a median white band followed by a dark-brown band. Body whorl averaging 60% of shell length, sculptured as spire; coloration pattern showing a subsutural light-brown band followed by a marked, white, narrow band halfway between the suture and the aperture, a wide, dark-brown band extending just below the posterior apertural angle, a whitish to brownish band as wide as the previous one gradually merging with the dark-brown, anteriormost band that extends to the anterior tip of the aperture. Aperture oval- elongated, rounded anteriorly, acute posteriorly; inner lip with three subequidistant teeth: one oblique colu- mellar tooth faintly continuous with a callus that runs along the anterior border of the aperture and connects to the tip of the outer lip tooth; two parietal teeth, the anterior one half the length of the aperture, perpendic- ular to the columellar axis and continuing inward; the posterior one usually strongest, not continuing inward; outer lip sharp, with a ridge-like tooth about opposite the anterior parietal tooth, penetrating inside the aper- ture for about half a whorl; a wide callus develops an- terior to the outer lip tooth. Protoconch 235x155 ^m, whitish, smooth, somewhat inflated, barely over one whorl, with elongated, narrowing lip (Fig. 21). Radula: (Figs. 31-32) Formula [31-h24-H-l-24-l-31] X 100-1-. Central tooth slightly above the plane of the lateral teeth; base narrow, subquadrangular, deeply emarginate with medial lateral projections flaring out; crown less than half the length of the lateral teeth, nar- row, unicuspid: mesocone somewhat sharp. Lateral teeth 24, base elongated, slightly bent medially, with lateral projections flaring out asymmetrically, the inner projec- tion smaller, at about the first third of the base, the outer one at about mid-length of the base; crown covering three quarters of base, quadrangular, with a longitudinal de- pression in the middle. Marginal teeth 31; basis barely longer than crown, diminishing gradually to completely disappear under crown, developing a basal ectocone; crown bicuspid, endocone sharp, short on first teeth, gradually elongating towards the end of the row, me- socone long and narrow, blunt at tip. Page 4 THE NAUTILUS, Vol. 109, No. 1 A. M. de Frias Martins, 199.5 Page 5 Remarks: The Hawaiian Allochroa bronnii is the largest species of the genus, sometimes reaching 12 mm in length (Souleyet, 1852; USNM 131288). It has been confused with larger forms of the highly variable Allochroa lay- ardi, an Indo-Pacific species (Dautzenberg & Fischer, 1905). They can be set apart by the narrower base of the aperture in A. bronnii, whereas shells of identical size of A. latjardi have a wider apertural base (see Figs. 1 and 5). In Hawaii, where both species overlap, they are morphologically very distinct (see Figs. 1, 2, 4). Although size alone seems to be sufficiently diagnostic to separate these two species, A. bronnii shows also a proportionally smaller number of whorls for the length of the shell. However, anatomical studies are needed to ascertain bet- ter the differences between these two closely related spe- cies. Habitat Notes: "Found seaward of Melampits, Pedipes, and Laemodonta. often buried in gravel in high tide pools and along rocky shores" (Kay, 1979). Range: Allochroa bronnii is endemic to the Hawaiian Islands. Material Examined: (Sample size in parenthesis). HA- WAIIAN ISLANDS: [USNM 20795 (5), 31582 (4), 98175 (6), 98176 (9), 98178 (3), 131288 (1), 319528 (4). 346615 (11)]; HAWAII ISLAND: Keei, near Napoopoo, Kona Coast [USNM 767596 (1)]; OAHU ISLAND: Honolulu [USNM 159574 (1)]; Pearl City, Honolulu [MCZ 55989 (1)]; 1935 dredgings, Hickham Air Force Base, Pearl Harbor [USNM 484416 (3)]; Kaneohe Bay [USNM 887316 (8)]. Allochroa latjardi (H. & A. Adams, 1855) Figures 3-14, 22-27, 30, 33-35, 39-45 Melampus layardi 'Ads.' Pfeiffer, 1854:147. Nomen nudum^ Ophicardcltis (Lainwdonta) latjardi H & A Adams, 1855a: 35 [Ceylon, lectotvpe herein selected BMNH 196980/1 (Fig. 3); two paralectotypes BMNH 196980/2-3]. Lai- modonla layardi (H. & A Adams) H. & A. Adams, 18.55b 246. Lainwdonta pfeifferi Dunker in Pfeiffer I860: 201 [Japan; location of type unknown); Dunker, 1861:25, pi 2, fig. 19. Lainwdonta conica Pease 1863: 242 [Pacific Islands; lectotype selected by Kay (1965) BMNH 1964292 (Fig. 4); five paralectotypes BMNH 1964293]. Laimadonta conica Pease. Pease, 1868101, pi 12, fig 15 (misspelling of Laimodonta). Lainwdonta anaaensis Mousson, 1869: 63 (Anaa Island, Paumotu group; location of type un- known]. Lacmodo7ita conica (Pease) Martens & Lang- kavel, 1871:57, Ptecotrema anaaensis (Mousson). Paetel, 1873:114. Lainwdonta hronii var. conica (Pease). Jickeli, 1874:178, pi. 7, fig, 3. Melampus pfeifferi (Dunker). Pfeif- fer, 1876:319. Melampus co7iicus (Pease). Pfeiffer, 1876: 319. PMelampus anaaensis (Mousson). Pfeiffer, 1876320. Marimtla (Laimodonta) conica (Pease). Nevill, 1878:220. Marinula (Laenwdonta) conica (Pease). Martens, 1880: 208. Laimodonta conica var conicoides Tapparone Ca- nefri, 1883:240 [Aru Island, New Guinea; location of type unknown]. Laenwdonta (Tralia) layardi (H. & A. Adams). Crosse, 1894:321. Melampus (Laimodonta) conicus (Pease). Kohelt, 1898:224, pi. 26, fig. 14. Melampus (Laimodonta) anaaensis (Mousson). Kobelt, 1898:228, pi. 27, figs. 10-11. Laimodonta bronni var. producta Dautzenberg & H. Fi- scher, 1905:108, pi. IV, figs 20, 21 [Anse du Crane, Bah'i'e d'Along, Tonkin; syntypes MNHN (Fig, 5)]. Enterodonta conica (Pease), Germain, 1921:264. Allochroa conica (Pease), van Benthem Jutting, 1941:283. Allochroa pfeif- feri (Dunker) Kuroda & Habe, 1952:38. Allochroa layardi (H. & A Adams) Franc, 19.54:365. Description: Shell (Figs. 3-14, 22-27) length to 8 mm, ovate-conical, fragile to somewhat solid; chestnut-brown to pale-yellow, banded. Spire moderately high, whorls 6.0 to 6.5, plano-convex, with spiral striae crossed by irregularly marked longitudinal growth lines which give the surface a faint granulose appearance. Body whorl about 70% shell length, sculptured as the previous whorls; color pattern showing a faint subsutural light band, a marked whitish band halfway between the suture and the aperture, also visible on the two previous whorls, sometimes with a third, wider pale-yellow band gradu- ally becoming dark-brown anteriorly. Aperture auricu- late; inner lip tridentate; one oblique columellar tooth not reaching the border of the lip; two parietal teeth, anterior lamellar, transverse, continuing inwards, pos- terior largest, thick and bifid in larger individuals, slight- ly elongated but not continuing inwards; outer lip sharp, with a ridge-like tooth about halfway along the length of the aperture, penetrating deep inside the aperture; a wide callus may develop anteriorly, as a continuation of the outer lip tooth, also penetrating inwards; a second outer lip tooth is sometimes present over the callus, lo- cated at a level between the anterior parietal tooth and the columellar tooth. Juveniles pilose, the hairs distrib- uted along the spiral striae (Fig. 12). Protoconch 210- 235 X 150-175 nm whitish, smooth, somewhat inflated, a little over one whorl, with rounded peristome (Figs. 22-27). Figures 1-2. Allochroa bronnii (Philippi); 1. Hawaii, USNM 98175, si = 8 2 mm; 2. Pearl City, Honolulu, Hawaii, MCZ 55989, si = 3.5 mm. Figures 3-12. Allochroa layardi (H. & A Adams);3. Lectotype, Ceylon, BMNH 196980.a, si = 7.0 mm, 4. Lectotype of Laimodonta conica Pease, Pacific Islands [Hawaii], BMNH 1964292, si = 4.2 mm; 5. Holotype of Laimodonta bronni var, producta Dautzenberg & Fischer, Bah'ie d'Along, Anse du Oane, Tonkin, China, MNHN (not numbered), si = 8 0 mm; 6. Paumotu [Tuamotu] Islands, specimen illustrated in Pease (1868, pi, 12, fig. 15), ANSP 22610 [type written in label], si = 7 0 mm; 7. Mataira Island, Raroia, Tuamotu Islands, USNM 698771, si = 5 8 mm; 8. North Beach, Henderson Island, Pitcairn Islands, R C. Preece coll , si = 6.1 mm; 9. Lijeron Island, Jaluit Atoll, Marshall Islands, USNM 6600.35, si = 4 6 mm; 10. Lugbon Island, Philippines, USNM 201147, si = .5.5 mm; 11. Riu Kiu Islands, Japan, USNM 345877, si = 5.0 mm; 12. Hirado, Hizen, Japan, USNM :345878, si = 4.1 mm. si = shell length. Page 6 THE NAUTILUS, Vol. 109, No. 1 Figures 13-14. Allorhroa layardi (H & A. Adams), Mataira Island, Raroia, Tuamotu Islands, USNM 698771, si = 3 5, and 2.5 mm. Figures 15-18. Allochroa nana Martins, n. sp., Eninman Island, Bikini, Marshall Islands; 15, 17-18. paratypes, USNM 585963, si = 2.2, 3.7, and 3.0 mm; 16. holotype, USNM 585963.a, si = 3.2 mm Figures 19-20. Allochroa tenuis Martins, n sp , Noumea, New Caledonia; 19. holotype, USNM 724374.a, si = 4.1 mm; 20. paratype, USNM 724374, si = 3.5 mm. si = shell length. Radula: (Figs. 33-35) formula [26 + 23+1+23 + 26] x 115. Centra! tooth slightly above the plane of the lateral teeth; base deeply emarginate, with lateral projections flaring out at mid-length, gradually tapering to neck region; crown half-length of that of lateral teeth, narrow, unicuspid; mesocono somewhat sharp. Lateral teeth 23; base elongated, slightly bent medially; a rounded inner prominence arises on the lower third, an outer projection flares out at mid-length and tapers towards neck; crown covering two thirds of base and as wide as the latter, unicuspid; cusp rounded. Marginal teeth 26; bases of first teeth barely longer than crown, gradually tliniinishing to completely disappear underneath crown, developing a conspicuous lateral basal cusp on the last 13 teeth; crown bicuspid; endocone, sharp, short on first teeth, gradually elongating to about three quarters of meso- cone; mesocone as long as that of last lateral teeth, round- ed at tip. Visceral mass: Conical, not following external winding of shell, with whorl impressions weakly insinuated, in- dicating the extensive resorption of the iiuier whorls of the shell. Digestive system: Mandible (Fig. 39) with fibrous ap- pearance, tripartite; median portion roughly quadran- A. M. de Frias Martins, 1995 Page 7 gular with extremities abruptly tapering posteriorly, lat- eral portions of sparse, loose fibers, tapering on both extremities. Salivary glands long, cylindrical. Stomach (Fig. 40) tripartite; cardiac region dilated, thin, receiving at its base the duct of anterior diverticulum; mid-portion comprising most of the stomach's volume, muscularized; gastric caecum thin, pouch-like, receiving posterior di- verticulum; pyloric region small, thin. Reproductive system: (Fig. 30, 41-44) Ovotestis of long, inverted claviform acini embedded in the conical posterior lobe of the digestive gland, connecting with a collecting duct by their thinner ends, the broader, dark- red dotted extremities appearing at the surface of the digestive gland as an irregular, winding band; proximal half of the hermaphroditic duct thin, relatively straight, distal half forming a greatly convoluted and somewhat dilated seminal vesicle; pallial gonoduct hermaphroditic to the vaginal aperture; albumen gland beneath the an- terior lobe of digestive gland and intestinal convolutions; posterior mucus gland tightly convoluted; anterior mucus gland and prostate gland covering entire length of pallial spermoviduct; bursa spherical, fitting under posterior mucus gland; bursa duct nearly equal in length to sper- moviduct, emptying near vaginal opening. Pallial gland long, sharply bent about midlength, anterior portion more dilated and slightly longer than posterior one, emptying at the female genital opening. Penial complex relatively short, proximal third dilated; vas deferens adhering to penial sheath, entering penial structure at the distal end of the proximal quarter; penis oval-oblong, blunt, con- stituted of a thickened, sharply bent, compactly folded muscular pilaster, lodged in the thin, membranous di- lation of the penial sheath; internal wall of the penial sheath with fine, radiating grooves at the site of the dilation, with two irregularly grooved longitudinal folds running forward to the aperture, one of them connecting distally to the penis, the other originating at the level of the tip of the penial papilla; penial retractor about as long as the penial complex, relatively thick. Nervous stjstem: (Fig. 45) Cerebral commissure short, about half the width of a cerebral ganglion; left cerebro- pleural connective twice the length of the right one; right cerebro-pedal connective somewhat longer than the left one; pleural ganglia small; right pleuro-parietal connec- tive very short, about onethird of the left one; left parietal ganglion as small as pleural ganglia, right one three times larger, with a thick pneumostomal nerve from where a rudimentary osphradial ganglion buds off; parieto-vis- ceral connectives relatively long, the left one somewhat longer. Remarks: Allochroa layardi was described by H. & A. Adams (1855a) from specimens from Ceylon [Sri Lanka], in the Cuming collection. H. & A. Adams' manuscript, although submitted for publication in 1854, was not pub- lished until January 1855 (see Duncan, 1937; Trew, 1992). Pfeiffer, who had access to the manuscript of the Adams brothers, listed their new species in his Synopsis (1854), giving credit to the original author(s) and placing the species under the genus Melampus. Pfeiffer, however. did not add any descriptive notes; for this reason, the name "Melampus layardi 'Ad.' Pfeiffer' should be treat- ed as nomen nudum. The Adams brothers' name was apparently ignored in the various subsequent descrip- tions of taxa here considered as synonyms of A. layardi, and comparisons referred rather to A. hronnii. Such was the case of Laimodonta pfeifferi. from Japan, and of Laimodonta Inonni var. producta from China (see Fig. 5). Allochroa laijardi, however, has a more slender over- all shape and is smaller than A. bronnii, mainly in Hawaii where both species coexist. Pease (1863) described Lai- modonta conica from the Hawaiian Islands (Fig. 4); five years later, however. Pease (1868) figured his Hawaiian species using a specimen from Paumotus [Tuamotus Is- lands], almost twice the size and more elongated than the type specimens (Fig. 6). Mousson (1869) described and figured the larger Tuamotu shells from Anaa Island as Laemodonta anaaensis. commenting on their clear relationship with A. bronnii, but at the same time calling attention to the smaller size of his species relative to the latter; his measurements (8 mm), however, are the largest known for A. layardi. Very variable across the Indo-Pacific, A. laijardi is usually markedly striated, although some specimens can be extremely smooth (Fig. 8). It also varies in shell thick- ness, the thick specimens sometimes showing an addi- tional internal lamella on the outer lip (Figs. 7, 9, 13), which Tapparone Canefri (1883) considered to be the variety conicoides of Laimodonta conica [=A. layardi]. The color pattern remains basically the same throughout; however, the banding delimitation may sometimes lack sharpness (Fig. 10). The shape and size of the protoconch are consistently homogeneous throughout the distribu- tional range of the species (Figs. 22-27). Allochroa bron- nii (Fig. 21) possesses a protoconch similar to that of A. layardi, also stressing the close affinities of the two spe- cies. However, as pointed out before, the size and ratio between the number of whorls and the length of the shell justify their taxonomic separation. Only one radula of each species was studied. It appears that the base of the central tooth of A. layardi is decidedly triangular (Fig. 33), whereas that of A. bronnii is nar- rowly quadrangular (Fig. 31) The crown of the lateral teeth of A. bronnii seems to be narrower and longer than that of A. layardi. Habitat Notes: Gassies (1863) stated that this animal ap- pears to be completely marine, and that Montrouzier found it under stones near the shore, together with col- umbellids, ranellids and tritons. Preece (personal com- munication) found them in crevices of exposed rocks at about high tide level. Range: Allochroa layardi is widely distributed through- out the Indo-Pacific; it has been reported from Mauritius (Morelet, 1882), Sri Lanka, India (Nevill, 1878), China, Japan, Philippines, New Guinea, Australia, New Cale- donia, Marshall Islands, Tuamotu Islands, Hawaii. Material Examined: (Sample size in parenthesis) CEY- LON [SRI LANKA] [BMNH 196980/1 (lectotype), Pages THE NAUTILUS, Vol. 109, No. 1 196980/2-3 (paralectotypes of Ophicardelus (Laimo- donta)layardi)]. CHINA; Bahie d'Along, Ansedu Crane, Tonquin [MNHN, not numbered, holotype and paratype of Laimodonta bronni var. producta]. JAPAN: Hirado, Hizen [USNM 345878 (30)]; Riu Kiu Islands [USNM 345877 (56)]; Loo Choo Islands [USNM 30811 (12), 161507 (2)]; Nago, Okinawa Island, Loo Choo Islands [USNM 363720 (1)]. PHILIPPINE ISLANDS: Bohol Island [USNM 302973 (4)]; off Point Origon, Tablas [USNM 281503 (1)]; Liigbon Island [USNM 201147 (7)]; Point Hinayangan, Leyte [USNM 887317 ex Chicago Academy of Sciences (3)]; Sogod Bay, Leyte [USNM 283799 (1)]; off Point Dumurug, Masbate [USNM 274834 (1)]; off Adyagan Island, E Masbate [USNM 281808(1)]; Ragay Gulf, off Arena Point, Luzon [USNM 284250 (1)]; off Matacot Point, W Louzon [USNM 259939 (1)]; off Pes- cador Island, Tanon Station [USNM 281169 (1)]; off Cap- itancillo Island, N Cebu [USNM 287800 (1)]; off Lanis Point, E Cebu [USNM 290562 (2)]; Pujada Bay, E Min- danao [USNM 255647 (1), 276398 (2)]; Tilig, Lubang Island [USNM 229914 (1)]. NEW CALEDONIA: Lifu [USNM 422639 (1), 422640 (1), 423167 (4)]. AUSTRA- LIA: [USNM 887318 (1)]; mouth of Bigota Creek, Barrow Island [USNM 691788 (1)]. COOK ISLANDS: Tupapa, E of Avarua, Rarotonga [USNM 721509 (1)]. MAR- SHALL ISLANDS: N end of Lijeron Island, Jaluit Atoll [USNM 660035 (8)]. TUAMOTU ISLANDS: [USNM 98177 (3), 711465 (6)]; RAROIA ISLANDS: Ngarumaoa Island [USNM 697524 (1), 697767 (1), 698079 (2), 720349 (17)]; S end of Ngarumaoa Island [USNM 697593 (3)]; Mataira Island [USNM 698771 (8), 711673 (1)]; Kumek- umo Island [USNM 698492 (5)]; Teteu Island [USNM 722685 (1), 722687 (1)]; S end of Teteu Island [USNM 722756 (50)]; Havana Island [USNM (1 fragment)]; Okaea Island [USNM 722834 ( 1 )]; S end of Opakea Island [USNM 722554 (79), 723548 (1)]; Oneroa Island [USNM 720768 (2)]; Ohava Island [USNM 720503 (19)]; Otikaheru Island [USNM 720684 (1)]; Vahi Tahi [USNM 613207 (1)]; Te- mao Harbor, Makatea [USNM 629788 (1)]; NE side of Tepukamaruia Island, Takumo[USNM 723724 (1)]; Motu Taraururoa, Mangareva Island [USNM 638224 (4)]. PIT- CAIRN ISLANDS: E end of North Beach, Henderson Island [R. C. Preece collection (4)]. HAWAIIAN IS- LANDS: [BMNH 1964292 (lectotype), 1964293 (5 para- lectotypes)]; USNM 15157 (2), 20796 (3), 20797 (12), 887319 (4)]. Allochroa nana Martins, new species Figures 15-18, 28 Description: Shell (Figs. 15-18, 28) length to 3.7 mm, ovato-conical, fragile, spirally striated, dark-brown to golden-brown, banded. Spire moderately high, 5 plano- convex, dark-brown to brown whorls, sometimes with a faint equatorial lighter band; spiral striae undulated due to crossing by marked growth lines which arc predom- inant on the early whorls of teleoconch. Body whorl about 70% of shell length, sculptured as previous whorls; band- ed color pattern as follows: a light-brown, fine band from the suture to the first spiral stria, a darker band on the next two striae, a wider, white band covering the next 3-4 striae, a dark-brown band extending to the posterior end of the aperture, frequently continuing to the anterior tip of the shell, sometimes broken up by a diffuse, lighter band extending from the posterior end of the aperture to about the level of the anterior parietal tooth. Aperture semicircular, oblique; inner lip equidistantly tridentate: one small, oblique columellar tooth, two parietal teeth, the anterior strongest and perpendicular to the colu- mellar axis, the posterior about as strong as columellar tooth; outer lip sharp, very rarely with an elongated thickening opposite the anterior parietal tooth and weak- ly projecting inside the aperture (Fig. 17). Juveniles pi- lose (Fig. 15). Protoconch 285x210 ^m, bulbous, light- brown, with growth lines showing as faint narrow bands, with elongated, somewhat widening lip (Fig. 28). Type Locality: Eninman Island, Bikini, Marshall Islands. From drift line along tide pool at N end of island, outer reef side. Material Examined: (Sample size in parenthesis) MAR- SHALL ISLANDS: Eninman Island, Bikini [USNM 585963 (holotype), 887320 (64 paratypes)]; Rojoa Island, Enewetak [USNM 587120 (1)]; Pujiyoru Island, Enew- etak [USNM 587101 (1)]; N end of Bock Island, Rongerik [USNM 594920 (2)J Etymology: nanus, i = dwarf. Range: Known from Bikini, Enewetak and Rongerik, in the Marshall Islands. Remarks: Allochroa nana is the smallest species of the genus, and size alone will easily distinguish it from the previously described species. It is, however, best char- acterized by its large protoconch with faint colabral bands marking the growth lines and with a somewhat widening lip (Fig. 28). Allochroa tenuis Martins, new species Figures 19-20, 29, 36-38 Description: Shell (Figs. 19-20, 29) length to 4.6 mm, ovato-conical, fragile, spirally striated, purple-brown to brown, banded. Spire moderately high, 5.5 plano-convex, spirally striated, dark-brown whorls, the last one darkest, usually purple-brown. Body whorl about 70% of shell length, sculptured as previous \\ horls; baniled color pat- tern as follows: a gradually darkening brown band ex- tending from the suture to the third stria, a marked white band over the next three striae, and a more or less ho- mogeneous brown region extending to the tip of the shell, sometimes broken up by a diffuse, lighter band between the end of the aperture and the posterior parietal tooth. Aperture ovate; inner lip equidistantly tridentate; one columellar tooth, obli(]ue, not reaching the border of the aperture lip, two parietal teeth, the anterior strongest and perpendicular to the columellar axis, the posterior about as strong as columellar tooth; outer lip sharp, rarely with an elongated thickening opposite the anterior pa- A. M. de Frias Martins, 1995 Page 9 Figures 21-29. Protoconchs of Allochroa species 21. Allochroa bronnii (Philippi), Hawaii, USNM 98175. 22-27. A. layardi (H. & A. Adams); 22. Hawaii, LISNM 20797; 23. paralectotype, Ceylon, BMNH 196980; 24. Henderson Island, Pitcairn Islands (R. C. Preece); 25. Lugbon Island, Philippines, USNM 201147; 26. Riu Kiu, Japan, USNM 345877; 27. Mataira Island, Raroia, Tuamotu Islands, USNM 698771 (specimen shown in fig. 14). 28. A. nana Martins, n. sp., Eninman Island, Bikini, Marshall Islands, lectotype, USNM 585963. 29. A. tenuis Martins, n. sp., Noumea, New Caledonia, lectotype, USNM 724347. Scale bar = 100 Mm. rietal tooth and weakly projecting inside the aperture (Figs. 19-20). Protoconch 385 x 240 ^m, bulbous, light- brown, showing 1.3 whorls, with somewhat reflected lip creating an umbilical slit (Fig. 29). Radula: (Figs. 36-38) Formula [17 + 14 + 1 + 14 + 17] X 100 + . Central tooth barely above the plane of the lateral teeth; base deeply emarginate, with lateral pro- jections Daring out at mid length; crown small, with nar- row, pointed mesocone. Lateral teeth 14, base elongated with lateral projections asymmetrical, the inner lower and serving as "point d'apui" for the articulation with the outer basal projection of the next tooth in the row (see Fig. 36); crown subquadrangular, long, with rounded mesocone. Marginal teeth 17, with one sharp endocone becoming longer towards the end of the row; mesocone becoming sharper towards the end of the row. Type Locality: Lighthouse island opposite Noumea, New Caledonia. From crevices on undersides of coral slabs, outer reef. Material Examined: (Sample size in parenthesis) NEW CALEDONIA: Lighthouse island opposite Noumea [USNM 724374 (holotype), 887321 (117 paratypes)]; N side of N'Go Bay [USNM 725042 (1)]; Touaourou [USNM 724628 (1), 724722 (1)]. PHILIPPINES: Pujada Bay, E Mindanao [USNM 887322 (1)]. Etymology: tenuis, e = thin. Range: Known from New Caledonia and Mindanao, Philippines. Remarks: Allochroa tenuis is readily distinguished from all other species in the genus on the basis of its thinner, darker shell and its large protoconch with flaring lip (see Fig. 29). The smaller, lighter but equally thin Allochroa nana, described above, also has a large protoconch, but without a flaring lip. The shape of the protoconch of A. tenuis, although unique among the known Allochroa, strongly resembles that of Ovatella aequalis (Lowe, 1832). Page 10 THE NAUTILUS, Vol. 109, No. 1 Figure 30. Internal structure of tlie peiiial comple.x of Alluchrua layardi (H. & A. Adams), Henderson Island, Pitcairn Islands Figures 31-38. Radulae of Allochroa species. 31-32. A. bronnii (Pfiilippi), Hawaii; 31. central tooth and first lateral teeth; 32. last lateral teeth and marginal teeth 33-35. A. layardi (H & A Adams), Henderson Island, Pitcairn Islands; 33, central tooth and first lateral teeth; 34. last lateral teeth and marginal teeth; 35, marginal teeth showing basal cusp (arrow ) 36-38. A tenuis Martins, n. sp,, Noumea, New Caledonia; 36, central tooth (arrow) and lateral teeth; 37. last lateral teeth and marginal teeth in a worn region; 38 last lateral teeth and marginal teeth in a fresh region. Scale bar = 100 fim for Fig. 30, 10 nm for all others. The relationships of Allochroa with this European genus are therefore strengthened. RELATIONSHIPS WITHIN THE ELLOBIIDAE Since the studies of Odhner (1925) and Morton (1955) the subfamilial division of the Ellobiidae has been based on anatomical characters, mainly on those of the repro- ductive system. Martins (in press a, b), studying the char- acteristics of the central nervous system, concurred with Morton s (1955) conclusions and provided a more com- plete subfamilial characterization. The Fythiinae, the most primitive subfamily, have a monaulic, entirely glan- dular, pallial gonoduct and a wide visceral nerve ring Figures 39-1-5. ,\natomy of Allochroa layardi (H, & A. Adams), Henderson Island, Pitcairn Islands, 39. mandible; 40. stomach; 41, 42. reproductive system; 43. penial complex; 44, internal structure of penial complex; 45. central nervous system. Scale bar = 2 mm for Figs. 41-42, I mm for all others. ad, anterior diverticulmn; agl, albumen gland; amgl, anterior mucus gland; avd, anterior vas deferens; br, bursa, brd, bursa duct; ca, stomachal caecum; car, cardiac region of stomach; eg, cerebral ganglion; fp, fertilization pouch; hd, hermaphroditic duct, mb, A. M. de Frias Martins, 1995 Page 11 muscular band; og, osphradial ganglion; ot, ovotestis; pd, posterior diverticulum; pdgl, posterior lobe of digestive gland; pe penis per, penial retractor muscle; pg, pedal ganglion; pgl, pallial gland, pil, penial pilasters; pig, pleural ganglion; pmgl, posterior mucus gland; pr, prostate gland/spermiduct; prg, parietal ganglion; psd, posterior vas deferens; pvl, pvloric region of stomach, sv, seminal vesicle; v, vagina; vg, visceral ganglion Page 12 THE NAUTILUS, Vol. 109, No. 1 witli a long right parieto-visceral connective. The Ello- biinae have a diaulic reproductive system with entirely glandular pallial gonoducts; the visceral nerve ring is wide and has a short right parieto-viscerai connective. The Carychiinae have a monaulic reproductive system with the prostate concentrated distally on the gonoduct and a central nervous system of the ellobiine type. The Pedipedinae possess a monaulic or incipient semi-diaulic pallia! gonoduct where only the pro.ximai half is glan- dular and a concentrated visceral nerve ring. The Me- lampodinae are characterized by an advanced semi- diaulic reproductive system with non-glandular pallial gonoducts and by concentration of the visceral nerve ring. Taking into account the preceding arrangement, Al- lochroa must remain within the Pythiinae in view of its monaulic, entirely glandular pallial gonoducts and the wide visceral nerve ring with a long right parieto-viscerai connective. Within the Pythiinae, Allochroa shows strongest affin- ities with the Mediterranean and Eastern North Atlantic Ovatella. based primarily on the presence of an internal lamella inside the outer lip, the protoconch of A. tenuis which, although different from all others in the genus, is very similar to that of Ovatella, and the configuration of the internal structure of the penial complex, where the penis appears as the development of one of the pi- lasters (Martins, in press b). ACKNOWLEDGMENTS I wish to express my gratitude to Dr. M. G. Harasewych, of the Division of Mollusks of the Smithsonian Institution, for his most valuable support. I also want to thank Dr. R. C. Preece, of Cambridge University, for providing me with preserved specimens. This work is integrated into the sabbatical project 1992/ 1993 and was partially funded by the Junta Nacional de Investigagao Cientifica e Tecnologica (JNICT). LITERATURE CITED Adams, H and A. Adams. 1855a [1854], Contributions to- wards the natural history of the Auricuiidae, a family of pulmoiiiferous Moliusca, with description of many new species from the Cumiiigian collection. Proceedings of the Zoological Society of Lontlon 22:30-37. Adams, H and A Adams. 1855b-1858 The genera of Recent Mollasca. J. van Voorst, London, 2:93-248, pis. 73-96[1855]; 285-412, pis. 97-112|185fij; 413-540, pis. 113-128 [1857]; 541-661, pis. 129-1.38(1858]. Ancey, C F. 1887 Nouveiles contributions malacologiques. IV: Auriculacecs d'Aden (Arable) 15iilletiii de la Societe Malacologiqiie P'ran^ai.se 4:273-299. van Bentheni Jutting, W. S. S. 1941. Non-Marine Moliusca from tlie satellite islands surrounding Java. Archives Neer- laiidaises de Zoologie 5(3):251-348 Crosse, H. 1894. Faune malacologique terrestre et fluviatilede la Nouvelle-Caledonie et de ses dependances. Journal de Conchyliologie 42:161-332; 333-473, pis. 7-10. Dautzenberg, P. and H. Fischer. 1905. 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Faune conchyliologique terrestre et fluvio- lacustrede la Nouvelle-Caledonie 1 e Partie J -B Bailliere et Fils, Paris, 126 pp., 8 pis. [Extract of "Actesde la Societe Linneenne de Bordeaux 24:211-330, 8 pis. "]. Germain, L. 1921, Faune malacologique terrestre et fluviatile des lies Mascareignes. F. Gaultier et A. Thebert, Angers, iv-l-495 pp., 13 pis. Gray, J E. 1855. List of Moliusca and Shells in the collection of the British Museum, collected and described by MM. Eydoux and Souleyet, in the "Voyage autour du Monde, " execute pendant les annees 1836 et 1837, sur la Corvette La Bonite, and in the "Histoire naturelle des Mollusques Pteropodes, " par MM. P.-C.-A.-L. Rang et Souleyet Print- ed by the order of the Trustees, London, 27 pp Gray, J, E. 1859. Systematic arrangement of the figures. In: Gray. M. E.. Figures of molluscous animals, selected from various authors, 2nd ed. Longman, Brown, Green and Longmans, Paternoster Row, London, l:pls. 1-78; 2:pls. 79-199;3:pls. 200-312; 4:iv-^219 pp.; 5:1-49 pp., pis. 313- 381. Jickeli, C. F. 1874. Fauna der Land- und Siisswasser-Molius- ken Nord-Ost-Afrika's. Nova Acta der Ksl. Leop.-Carol. Deutschen Akademie der Naturforscher 37(1): 1-352, 11 pis. Kay, E. A. 1965. Marine Molluscs in the Cuming collection, British Museum (Natural History) described by William Harper Pease Bulletin of the British Museum (Natural History), Zoology, Supplement 1:1-96, 14 pis. Kay, E. A. 1979. Hawaiian marine shells. Reef and Shore Fauna of Hawaii, Section 4: Moliusca. Bernice P. Bishop Museum Special Publication No. 64 (4). Bishop Museum Press, Honolulu, 653 pp., 195 figs Kobelt, W 1897-1901. Die F'amilie Auriculacea In: Martini & Chemnitz, Systematisches Ckmchylien-Cabinet Verlag von Bauer & Raspe (Emil Kiister), Niirnberg 1(16)1-76, pis. 10, 11, 13 [1897]; 77-228, pis. 12, 14-30 [1898]; 229- 268, pis. 31, 32 [1900]; 269-316, pi. 33 [1901]. Kuroda, T and T. Habe. 1952. Check list and bibliography of the Recent marine Moliusca of Japan Hosokawa Print- ing Company, Tokyo, 210 pp Martens, E. von. 1880 Molluscken /;i; Miibius, K. A., Bei- triige zur Meerestauna der Insel Mauritius und der Sey- chellen Verlag der (iutmann schen l?uchhanilluiig [Otto Enslin], Berlin, pp 181-352, pis 19-22 Martens, E. von and B Langkavel. 1871. Donum Bismarck- ianum. Eine Sammlung von Siidsee-Conchylien. Verlag von Ferdinand Berggold, Berlin, (2)-l-74 pp., 4 pis. A. M. de Frias Martins, 1995 Page 13 Martins, A M F In press a. Anatoiiiy and systematics of the western Atlantic Ellobiidae (Gastropoda, Pulmonata) Malacologia. Martins, A M F. In press b. Relationships within the Ello- biidae. Journal of Molluscan Studies Moreiet, A. 1882 Observations crititiues sur le Memoire de M. E. V. Martens, intitule: Mollusques des Mascareignes et des Sechelles Journal de Conchyliologie 30:85-106. Morton, J. E. 1955. The evolution of the Ellobiidae with a discussion on the origin of the Pulmonata. Proceedings of the Zoological Society of London 125:127-168. Mousson, A. 1869. Description de ciuelciues nouvelles especes terrestres de divers points de I'ocean Pacifique et de 1 Aus- Iralie. Journal de Conchyliologie 1755-68, pi. 5. Nevill, G. 1878. Handlist of Mollusca in the Indian Museum, Calcutta, Pt. I, Gastropoda Pulmonata ami Prosobranchia- Neuribranchia. Printed by order of the Trustees, Calcutta, 338 pp. Odhner, N. H 1925 Marinula juanensis. n. sp nebst Be- merkungen ijber der Systematik der Ellobiiden Arkiv for Zoologi, Stockholm I7A:1-15. Paetel, F. 1873. Catalog der Conchylien-Sammlung Verlag von Gebriider Paetel, Berlin, 172 pp Pease, W. H. 1863 [1862]. Description of new species of ma- rine shells from the Pacific Islands. Proceedings of the Zoological Society of London 16:240-243. Pease, W. H 1868. Descriptions of marine Gastropodae, in- habiting Polynesia. American Journal of Conchology 4: 71-80; 91-102, pi. 12. Pfeiffer, L, 1853. Studien zur Geschichte der Auriculaceen Zeitschrift fijr Malakozoologie 101-10 Pfeiffer, L 1854. Synopsis Auriculaceorum. Malakozoolo- gische Blatter 1: 145-1,56 Pfeiffer, L. 1856 Monographia Auriculaceorum viventium. T. Fischer, Cassel, 209 pp. Pfeiffer, L 1860. Zwei neue Auriculaceen. Malakozoologische BIStter 6:201-202. Pfeiffer, L. 1876. Monographia Pneunionopomorum viven- tium, accedenle fossilium enumeratione. Supplementum tertium, Monographiae Auriculaceorum parte secunda auctum T Fischer, (tassel, pp 241-479. Philippi, H A 1846 Diagnoses testaceorum quorundam no- vorum. Zeitschrift fiir Malakozoologie 3:97-98 Smith, B. J. 1992. Non-Marine Mollusca. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia. AGPS, Can- berra, vol. 8, xii-l-405 pp. Souleyet, L. F. A. 1852. In: Eydoux, F. & L F A. Souleyet. 1841-1852. Voyage autour du monde execute pendant les annees 1836 et 1837 sur la corvette La Bonite com- mandee par M Vaillant Zoologie Arthus Bertrand, Paris, 1(1) [I841]:iv-fxxxix + 1-134 pp.: 1(2) [1842]:135-334; 2 [1852]: 1-664. Souverbie, M. and R. P. Montrouzier 1864a. Descriptions despeces nouvelles de I'Archipel caledonien Journal de Conchyliologie, 12:41-42 Souverbie, M and P Montrouzier 1864b Descriptions des- peces nouvelles de I'Archipel caledonien Journal de Con- chyliologie, 12: 261-275, pi. 10 Tapparone tlanefri, C. 1883. Fauna Malacologica della Nuo- va Guinea e delle isole adiacenti Parte I. Molluschi Es- tramarini. Tipografia del R Istituto sordo-Muti, Genoa, 313 pp, II pis. Trew, A 1992. Henry and Arthur Adams's new molluscan names. National Museum of Wales, Cardiff, 63 pp. Zilch, A 1959 Gastropoda In: Schindewolf, O. H., Hand- buch der Palaozoologie, 6. Borntrager, Berlin, pp. 1-200, 701 figs. THE NAUTILUS 109(l):14-37, 1995 Page 14 A remarkable species richness of the Barleeidae (Gastropoda: Rissoacea) in the Eastern Atlantic Serge Gofas Museum National dlli.stoire Naturelle 55 rue Buffoii F-75005 Paris, France ABSTRACT The famiK Barleeidae, hitherto known in the Eastern Atlantic from tuo European and Northwest African species ol Barlecia and two species of Trupidurissuia from Sao Tome, is found to be represented by 18 species, 14 of which (S Barleeia, 3 Pseu- dodiala, 3 Lirobarleeia) are here described as new. Three new species share several shell or radular characters with the type species of Pseudodiala and not with Barleeia For this reason Pseudodiala is considered to he a valid genus. There are four centers of diversity; Senegal (3 species), the Cape Verde Islands (3 species), Sao Tome and Principe ((i species) and Southern Angola (6 species), together making the Eastern Atlantic the second most species-rich area after Western America, and the area with the highest diversity at the genus level. Ki'y words: Barleeidae, western Africa, new species, species diversity, biogeography INTRODUCTION The Barleeidae is a family of minute prosobranchs that typically live on intertidal or shallow subtidal hard bot- toms of temperate to tropical areas. Many species live within coralline algae, or on "maerl"', deposits of living calcareous algal concretions. The genera of the Barleei- dae have been reviewed by Ponder (1983), who then considered the Anabathriiiae as a subfamily of the Bar- leeidae but later raised tlieui to full family status (Ponder, 1988). Western America, from California to Panama and the Galapagos Islands, is the area with most described species of Barleeidae: 25 species are recognized as valid b\ Pon- der (1983). I'he fainily is also represented in the (krib- bean, eastern South America, East and South Africa, South-east Asia, Australia and New Zealand, and Japan, but generally with low (1-5) s()ecies richness. The family is known hitherto in the Eastern Atlantic by four species: the mainly European Barleeia unifas- ciata (Montagu, 1803), a less well known species B. goug- eti (Michaud, 1830) from NW Africa, and two species of Tropidorissoia from the Island of Sao Tome, in the Gulf of Guinea (Tomlin & Shackleford, 1915; Rolan & Templado, 1994). Several species that may be assigned to the Barleeidae were described from the island of St. Helena (Smith, 1890), but the systematic position of most of them remains uncertain until living animals are ex- amined. Recent collecting in Angola by the author, and other collectors' material from West Africa in Museum Na- tional d'Histoire Naturelle of Paris and Museo Nacional de Ciencias Naturales, Madrid, has shown that there are a number of undescribed species of Barleeidae on the mainland coast of West Africa, the Cape Verde Islands, Sao Tome and Principe, making this area one of the centers of species diversity for the family. No species assignable to the Anabathridae has been found in this material, with the exception of the southernmost records of the European species Nodiilus conturtus (Jeffreys, 1856) in Mauritania. The material examined has the following origins: — dredgings and shore collecting bv I. Marche-Mar- chad, 1953-1956, mostly from Senegal (MNHN). — intertidal collecting in the (>anarv Islands (1981) and Mauritania (1983) by P. Bouchet (MNHN). — the author's collecting in Angola and Sao Tome, 1981-1987, and in Europe (MNHN). — intertidal and SCUBA collecting in the Cape Verde Islands bv the 'la expedicion C^ientifica Iberica a Cabo Verde ", August 1985 (MNCN). — intertidal and shallow subtidal collecting by E. Ro- lan in Sao Tome, Principe and the Cape Verde Is. (MNCN and private collection of E. Rolan, Vigo). Radulae have been examined using SEM for a number oi species. Radular morphology has been found infor- mative for classification at the family and genus level, but not lor the disirimination of clo.seK related species. ABBREVIATIONS AMS = Australian Museimi, S\(lne\ CER = privati' collet'tion ol Emilio Rolan, V igo. MNHN = Museum National d'Histoire Naturelle, Paris. MNCN = Museo Nacional de CUencias Naturales, Madrid. UAN = Universidade Agostinho Neto, Luanda, cat. = catalogue number. S. Gofas, 1995 Page 15 leg. = "legit ", collected by. spec, specs. = live-collected specimen(s). sh. = sheil(s). jii\ . = juvenile(s). SEM = Scanning electron micrograph(s). * denotes specimens in the wet collection. SYSTEMATICS Family BARLEEIDAE Gray, 1857 The family is diagnosed by having: a pitted protoconch, a groove on the sole of the metapodium reaching its posterior end, and a single-layered operculum with an inner peg. This last character differentiates the Barleei- dae from the Anabathridae Pallial and metapodial ten- tacles are t\picalK absent. The reader is referred to Pon- der (1983) for a detailed account of the famii\. Genus Barlecia Clark, 1855 Type species: Turbo ruber Adams, 1797 (= Turbo twi- fasciatus Montagu, 1803), non Turbo ruber von Salis, 1793, by monotypy. Shell with smooth, solid teleoconch in all species consid- ered, non-umbilicate or very narrowly umbilicate, gen- erally with reddish tinge. Aperture ovate, slightly pro- socline to orthocline, with smooth lip thickened at some distance inside, and then beveled to a thin edge. Head- foot typically pigmented with black and yellow. Oper- culum with an internal peg, a smooth nucleus on the columellar side, and fine growth lines on the remainder of the external surface; a conspicuous external ridge along the columellar side, overrunning the nucleus. Radula (Ponder, 1983 and herein. Fig. 71-72): central tooth with a large rectangular median cusp and 2-3 small lateral denticles, one sharp basal denticle on each side, separated by a broad, hardly prominent lamella. Lateral teeth (one pair) with a rather broad base, terminating with a large, rather trapezoidal cusp flanked by 2-3 smaller cusps to the inner side and the outer side; with a U-shaped projection beneath those. Marginal teeth (two pairs) narrow, hook-shaped, with small, unipectinate cusps towards the distal end. Barleeia unifasciata (Montagu, 1803) (Figures 1, 29-30) Turhu ruber Adams, 179766, pi. 13 fig, 21-22 (iinii Tinlio niher von Sails, 179.3). Type locality; Pembrokeshire (Ikitisii Isle.s). Turbo unifasciatus Montagu, 1803:327-32H. T\pe locality; Southampton, and Island of Burrow, Devon (British Isles) Rissoa fulva Michaud, 18.30:12 (repr. 18.32:15). pi. 1. fig.l7- 18. Type locality: not specified; localities mentioned: Agde, Sete (Mediterranean France) and Corsica. Sabanaca hinf^hamiana Leach. 1852:1.54. Type localit\ ; Tor- bay, Devon (British Isles). Barlecia rubra var. clongata Bucquo) , Dautzenberg & Dollfus, 1884:316, pi 32, fig. 23. Barleeia clongata "nsp." Locard. 1886:272 and 576. Type localit) : Roussillon (Mediterra- nean France). Material examined: (collected by the author unless oth- erwise stated; all in MNHN) Europe: Guernsey, Vazon Bay, at low tide among corallines, 1000+ specs., 9.1994; Guernsey, Rocquaine Bay, 1000-h specs., 9.1994. Plou- manach (Finistere), at low tide on Corallina, 300 specs, (many juv); Lampaul-Ploudalmezeau (Finistere), on Corallina, 1000-1- specs, (mostly juv.; adults 2.3 x 1.3 to 2.8 X 1.8 mm); He de Sein (Finistere), 300 specs.; Guethary (Pyrenees-Atlantiques), 54* spec. (2.9 x 1.7 to 3.3 X 2.0 mm); St. Jean de Luz (Pyrenees Atlantiques), .30* -t- 150 specs. (2.4 x 1.4 to 3.2 x 1.8 mm); Hendaye (Pyrenees-Atlantiques), 200 specs.; San Sebastian (Gui- puzcoa), 1000-t- specs.; Ondarroa (Guipuzcoa), 1000-1- specs.; Calahonda (Malaga), 200 specs., Los Escullos (Al- meria), 7 specs. Strait of Gibraltar, Ceuta: Benzu, low tide, 4 specs. Morocco: Punta Cires, intertidal in algae, 100 specs.; Tangier Ba>, El Ghandouri, 20 specs.; Rabat, near Lahlou, 12 specs., Mohammedia (=Fedala), 30 sh.; Near Tarfaya, .35 specs., Ortea leg. 8.1987 (2.5 x 1.5 to 2.7 X 1.7, pattern of foot checked on rehydrated spec- imen). Mauritania: Bale de I'Etoile, 250 specs. (1.8 x 1.2 to 2.5 X 1.5 mm), Bouchet leg. 1983; Cap Blanc, exposed rocky shore, 18 specs. (16 juv.), Bouchet leg. 1983; Pointe des Maures, 100-1- specs., Bouchet leg. 1983. Senegal: Dakar area, 500 sh., Marche-Marchad leg.; Anse Bernard, 5 specs., Marche-Marchad leg. Tenerife, Ca- nary Islands: Playa Santo Domingo, 75 specs. (50 juv.), Bouchet leg. 7.1981; Punta del Hidalgo, 200 specs. (30 juv), Bouchet leg. 7.1981; La Tejita, intertidal, .34 sh. (10 juv), Bouchet leg. 7.1981; Palm-Mar, intertidal, 25 specs., Bouchet leg. 1981. Description: Shell conical, solid, adults 2.3 x 1.3 to 3.3 X 2.0 mm. Protoconch dome-shaped (450-500 /u in di- ameter) of 1 '/2 to 1% whorls, with very minute spiral rows of pits. Teleoconch of 3'/4 to 3'/2 whorls. Spire whorls rather flat, body whorl rounded and imperforate. Outer lip slightly prosocline, beveled inside to a thin edge. Color of shell reddish brown, paler towards the parietal inser- tion of the body whorl; plain or with whitish spiral bands, subsutural, median on body whorl and/or periumbilical; rarely entirely of a pale, wax-like color. Operculum dark crimson. Head-foot vsith superficial plain black pattern on upper part of head, snout, sides of foot and opercular lobes, upper part of propodium with black area extend- ing on the edges, and grading to a broad axial white area; sole entirely colorless. Yellow axial bar on tentacles, large, triangular yellow granular masses behind each eye. Habitat: On rocky shores, among red algae (preferably Corallina) at low water mark or in tidal pools (Southgate, 1982; Borja, 1986a, 1986b, 1987); in shallow photophilous algal mats in the Mediterranean. Distribution: From the British Isles to Senegal, the Ca- nar\ Is. and in the Mediterranean (including the Adriatic and the Eastern Mediterranean, but not the Black Sea). Page 16 THE NAUTILUS, Vol. 109, No. 1 Fifiurc I. litirlccia utiifasciata (Montagu, 1803), living animal from Oiita Length of sfiell 2.6 mm. Figure 2-3. Barlccia gougcti (Midiaud, IS.'iOj. 2. Living animal from Outa. Lt-ngth of .shell -4. 3 mm. 3. Specimen from Dakar, length 5.6 mm. Remarks: The above description is based on specimens from Western Europe and Morocco; such typical B. uni- fasciata are found as far south as Tarfaya, Morocco. The populations examined from Haie de I'Etoile, Mauritania and from Senegal are smaller, paler, with more variable prohle, but are tentatively assigned to B unijasciala pending further study. Bellon-Humbert (1974) has figured Barleeia iinifas- ciata from Morocco as "Barleeia niintiscula Montero- sato , and the sympatric B. guugeli as "B. rubra" , Mon- terosato's (1889) description and indication of size (1.5 mm) for B. minuscula, although laconic, fit Coriandria ftilgida (.Adams, 1797), a common species on the entire Atlantic coast of Morocco I consider Barleeia inirjuseiila a syiionvin ot Coriandria julgida, a view further .sup- ported 1)\ the fact that Monterosato lists both B. rubra aiul B. gnugeti but no species referable to Coriandria. Populations from the (]anarv Islands cannot be distin- S. Gofas, 1995 Page 17 guislied concliologicall\ trom tliose of Western Europe, and are thus treated as conspecific although it is likely that very little, if any, genetic exchange occurs. The head-foot of C^anarian specimens is darker, and the entire propodiuin is black, Barleeia gougcti (Michaud, 1830) (Figures 2-3, 31-32) Hissoa gougcti MichaiKl, 1830 9, pl.l. fig 7-S T\ [)e l(ic,ilit\ Senegal Hijdrohia ciwipacta Jeffress, 1883:120, pl.9. tig.9. (non Hij- drohia compacta Carpt-iiter, 1S64). Type lncalit\ : Taiigiers (Morocco), ? Barleeia majuscula Montfni.sato, 1884:69, T>pe locality: not specilied; localities mentioned: Ognina (Sicily), Alger (Al- geria), Lanipedusa (Sicily channel), Barleeia gnugeti var, unicnhir Montero.sato. 1889:34 (Casa- blanca, Morocco) Barleeia gougcti var, ccrca Monlcrosato, 1889:34 (t^asablanca, Morocco), Barleeia carrozzai van Aartsen & Giannuzzi-Savelli, 1991 (re- placement name for Hydrobia compacta). Material examined: (collected by the author unless oth- erwise stated; all in MNHN) Spain: Tarifa, intertidal, 3 specs. (3.5 X 2,1 mm). Strait of Gibraltar, Ceuta: Punta Almina 35-40 m, 38* -H 12 specs. (4.3 x 2.3 to 5.5 x 2.8 mm), Bouchet leg. 5.1986; Benzii, 8 sh. (white). Mo- rocco: Temara, low tide, 10 spec; Mouth of Oued Yquem (= Rose-Marie), S. of Rabat, 31 spec, de Lepinev leg.; Mohammedia (= Fedala) 50 sh.; El Jadida (= Mazagan), low tide, 400 specs.; Essaouira (= Mogador), 300 specs. (2.7 X 1.6 to 6.1 X 3.0 mm); Near Tarfaya, 12 specs., Ortea leg. 8.1987. Mauritania: Bale de I'Etoile, 140 specs. (2.9 X 1.7 to 4.3 X 2.3 mm), Bouchet leg. 1983. Senegal: Dakar, 8 specs. (5.0 x 3.1 to 5.9 x 3.2 mm), Nickles leg. 1946; Goree, 7 sh. (pale periumbilical zone), H. Fi- scher collection; Goree, bale de St. Jean, 50 specs., Delais leg. 1.1962; Goree 20 m, 10 specs.. Pin leg 1994; Bel- Air near Dakar, 6-8 m, 59 specs. (3.1 x 1.8 to 5.0 x 2.7 mm, with attached Crepidula), Marche-Marchad leg.; Dakar, Anse Bernard, 9 specs., Marche-Marchad leg.; Dakar-ville, 10 specs.. Pin leg. 1994. Description: Shell conical, solid, adults 2.7 x 1.6 to 6.1 X 3.0 mm. Protoconch dome-shaped (500 n in diameter), of l'/2 to 1% whorls, with very minute spiral rows of pits, Teleoconch of 4 to 5 whorls. Spire whorls rather flat, profile of body whorl very faintly carinate and imper- forate. Outer lip strongly prosocline near its parietal in- sertion, then nearly orthocline. Columella protruding an- teriorly, more pronounced in juveniles, onl\ slightK in adults v\ here the aperture is more rounded. C^olor of shell reddish brown, paler towards the parietal insertion of the body whorl; plain or with periumbilical whitish spiral zone; sometimes entirely of a pale, wax-like color. Oper- culum dark crimson. Head-foot with superficial plain black pattern on upper part of head, snout, and opercular lobes; upper part of propodium and sole entirely color- less. Yellow axial bar on tentacles, large, yellow granular masses behind each eve and (most often) in each oper- cular lobe anteriorly to the black markings. Habitat: On hard bottoms with algal concretions, low intertidal to 40 m. Distribution: From the Strait of Gibraltar to Senegal. Remarks; This large species occurs sympatrically with B unifasciata all along the coast of NW Africa, from the Strait of Gibraltar .southwards. Barleeia gougcti is usualK subtidal and is restricted to the lowermost part of the tidal zone, whereas B. unifasciata is essentially an intertidal species. The relative abundance of both species is variable, with B. gougeti the most abundant species south of Rabat in Morocco. The ranges of sizes in the two species overlap, and the boundary cannot be clearly placed with onl\ concholog- ical characters. The most obvious distinctive character is the pigmentation pattern of the propodium, which has lateral black areas in B. unifasciata (a non-variable fea- ture in Brittany and the Basque country where 6. uni- fasciata occurs alone), and is white in B. gougeti. The columella of B. gougeti is more protruding than in B. rubra. Populations where an entireK pale shell is the dominant pattern are common in B. gougeti. Barleeia verdcnsis Gofas, new species (Figures 4, 33-34) Type material: Holotype and 120 paratypes (MNCN cat. 15.05/20526); 120 paratypes (MNHN) collected alive from the type locality. Type locality: Salmanza, Sao Vicente, Cape N'erde Is- lands, (16°54'N,24°57'W). Material examined: (collected by E. Rolan except spec- imens with date 8.1985 collected by "la expedicion ib- erica , or where otherwise stated) Cape Verde Islands, Boavista: Derrubado, 13 juv. sh. (CER); Rife de Chaves 4 m, 4 sh.+ 25* specs. 5.1988 (CER); Baia Teodora 2 m,32 + 12*specs. 4. 1988 (CER); Morro da Areia, 5.1986 (MNCN); Sal Rei, 40 sh., 5.1986 (MNCN) and 38 sh. (CER). Brava: Furna, 8 sh. (CER). Sal: Mordeira, 3 specs., 5.1986 (MNCN) and 20 sh. (CER); Fontona, 125 specs. (mostly juv), 8. 1985 (MNCN); Curral do Dado, 14 specs. (10 juv.) (MNCN); Joaquin Petinha, 157 specs., 8.1985 (MNCN); Parda, 1 spec. (MNCN); Palhona, 40 sh. + 4* specs., 5.1987 (CER); Sal (without further precision), 44 sh. (CER); Serra Negra, 4* specs., 5. 1987 (CER); Calheta Fonda 2-5 m, 15* specs., 5.1988 (CER); Palmeira, 2 + 65* specs, 8.1985 (CER); Rigona, 7 specs., 8.1985 (MNCN). Sao Nicolau: Tarrafal, 3 sh., 8.1985 (MNCN). Santiago: Tarrafal, 38 sh. (CER). Sao Vicente: Salmanza, intertidal, 10 specs., Cosel leg. 12.1978 (MNHN); the type material (2.7 x 1.6 to 3.3 x 1.8 mm) and 100-t- specs., 8.1985 (MNCN); 12 specs., 5.1987 (CER); Calhau, 4 sh. (CER). Description: Shell conical with rather tall spire, very solid, adults 2.7 x 1.6 to 3.3 x 1.8 mm (holotype 3.2 x Page 18 THE NAUTILUS, Vol. 109, No. 1 8 Figure 4. liarlceia verdensis Gofas, new .species, liolotype from Salmanza (Sao Vicente, Cape Verde I,s.), length .3 2 mm Figure 5. Barleeia aemilii Gofas, new species, liolotype from .Saragasa (.Sao Vicente, C:ape Verde Is.), length 1.7 mm Figure 6. Barlccta chefiae Gofas, new species, liolotype from Hife de Chaves (Sal, C:ape Verde Is.), length l.;5 mm Figures 7-8. Barleeia lumensis Gofas, new species. 7. Holotype from I'raia das Conchas (Sao Tome), length 2.1 mm 8. Living specimen from Praia das Con- chas (Sao Tome), length 2 1 mm. Figure 9. Barleeia laeniolata (Jofas, new species, iiolot\pe from Santa Ana (Sao Tome), length 1.6 mm. S. Gofas, 1995 Page 19 1.8 mm). Protoconch dome-shaped (.'350-40U n in di- ameter), ol 1 '/i whorls, with very minute pits arranged in crowded spiral rows. Teleoconch with 4 whorls. Spire whorls rather Hat, body whorl rounded, imperlorate. Outer lip slightly prosocline, beveled both outside and inside to a thin edge; parietal callus narrow. Color ot shell dark reddish brown, rather uniform except for a paler area towards the parietal insertion of the body whorl and a poorly defined paler bantl around the col- umellar area. Head-foot black except for the sole, the sides of the snout, and the tentacles. Yellow axial bars formed by crowded granules on tentacles; yellow gran- ular masses on opercular lobes. Habitat: Intertidal or shallow subtidal, among algae. Distribution: Cape Verde Islands. Remarks: This species resembles B. iinifasciata and has been identified as "B. rubra" in the literature (e.g. (^osel, 1982 and references therein). It is distinguished b\ its smaller protoconch, with more crowded rows ot pits, taller spire, and peculiar profile of the bod\ whorl, due to the fact that the aperture is beveled not only inside as in other Barleeia, but also outside. It is distinguished from B. goiigeti by its smaller size and less protruding columella. Barleeia aemilii Gofas, new species (Figures 5, 35-36) Type material: Holotvpe and 40 paratypes (MNCN cat. 15.05/20527); 40 paratypes (MNHN) collected alive from the type locality. Type locality: Saragasa, Sao Vicente, Cape Verde Islands (I6°49'N, 24°52'W). Material examined: (collected by E. Rolan, except spec- imens with date 8.1985 collected by "la expedicion ib- erica ) Cape Verde Islands, Boavisla: Sal Rei, 2 sh. (MNCN); Rife de Chaves, 4 m, 1 sh., 5. 1988 (CER). Santa Luzia: Praia Francisa, 3 sh. (CiER). Sal: (without precision) 3 sh. (CER); Joaquin Petinha, intertidal, 9 specs., 8.1985 (MNCN) and 1 spec. (CER); Mordeira, 1 sh., 8.1985 (CER ); Serra Negra, 1* spec, 5.1987 (CER); Palhona, 4 sh. (CER); Palmeira, on wharf, 8.1985, 2 specs. (MNCN) + 1* spec. (CER); Rigona, 1 spec, 8.1985 (MNCN); Rabo de Junco, 20 sh., (CER). Santiago: S of Tarrafal, 7 specs. + 16 sh. (CER); Prainha, Praia, 6 specs. (CER). Sao Vicente: Calhau, 11 sh. (CER); Saragasa, the type material, 8.1985 (1.4 x 0.85 to 1.75 x 1.0 mm); Salmanza, intertidal, 8.1985, 1 spec (MNCN) + 30 specs., 5.1987 (CER) + 4* specs. (CER); Mindelo, Baia Matiota, 1 spec, Cosel leg. 12.1978 (MNHN). Description: Shell conical, solid, adults 1.4 x 0.85 to 1.8 X 1.0 mm (holotype 1.7 x 0.95 mm). Protoconch dome- shaped with flattened top (280-300 ^ in diameter), l'/4 whorls, with very minute spiral rows of pits. Teleoconch of 3 to 3'/4 whorls. Spire whorls rather flat, body whorl very slightly angulated. Outer lip slightly prosocline, bev- eled inside to a thin edge; parietal callus narrow, de- tached over a tiny umbilical chink. Color of shell pale orange brown, paler towards the parietal insertion of the body whorl; with a sharply defined paler band along the angle of the body whorl, and a less clear pale band around the umbilical chink. Parietal callus tinged with brown. Shell of some individuals entireK of a pale, orange-brown color. Periostracum dull. Operculum reddish brown, more intense along the ridge and edges. Head-foot pale, with yellow axial bars on tentacles; greenish, iridescent buccal mass visible in the head b\ transparency. Habitat: Intertidal or shallow subtidal, among algae. Distribution: C-ape Verde Islands. Remarks: This species is readily distinguished from the previous one by its smaller size, banded color pattern and more pupoid profile. It is most closely related to B.chefiae n.sp , \\ hich is still smaller, has a more rounded periphery of the body whorl, and has a glossy periostra- cum rather than the dull, opaque texture found in B. aemilii. This species is dedicated to Dr. Emilio Rolan, of Vigo (Spain), a distinguished amateur who has been involved in most of the recent collecting in the Cape Verde Islands. Barleeia chefiae Gofas, new species (Figures 6, 37-38) Type material: Holotype (live collected) and 2 paratypes (sh.) from the type locality (MNCN cat. 15.05/20528), 12 parat\ pes (sh.) from Porto P>rreiro (MNCN cat. 15.05/ 20529), 24 paratypes (sh.) from Sal Rei (12 MNCN cat. 15.05/20530, 12 MNHN). Type locality: Rife de Chaves, Boavista, Cape Verde Islands, in 4 m. Material examined: (collected by E. Rolan except spec- imens with date 8.1985 collected by "la expedicion ib- erica ) Cape Verde Islands, Boavista: Derrubado, 2 sh. (CER); Baia Teodora, 1 sh. (CER); Porto Ferreiro, 4 m, 12 sh. (paratypes); Rife de Chaves, 4 m, the holotype, 2 paratypes (sh) and 6 sh., 5.1988; Sal Rei, 24 sh. (para- types), 8.1985. Brava: (no further details), 39 sh. (CER); Furna, 8 specs. (CER). Santa Luzia: Praia Francisa 1 m, 4 sh. (CER ). Santiago: Tarrafal, 2 sh. (CER). Sal: Pal- hona, 1* spec (CER). Sao Vicente: Saragasa, 2 specs., 8.1985 (MNCN); Pedrinha 2 sh. (CER). Description: Shell pupoid, moderately solid, translucent, adults 1.1 X 0.7 to 1.3 x 0.75 mm (holotype 1.3 x 0.75 mm). Protoconch dome-shaped with flattened top (260- 270 n in diameter), I'A whorls, with very minute spiral rows of pits. Teleoconch of 3 to 3Vi whorls. Spire whorls rather flat, body whorl rounded. Outer lip very slightly prosocline, beveled inside to a thin edge; parietal callus narrow, detached over a tiny umbilical chink. Color of shell brownish, with two sharply defined paler bands along the periphery of the body whorl, and around the Page 20 THE NAUTILUS, Vol. 109, No. 1 umbilical chink, and a broad, ill-defined pale area run- ning along tlie v\ liorls almost from suture to suture; a dark subsuturai line ("false suture ) corresponds to the inner surface of the whorl seen by transparency. Parietal callus and periumbilical area dark. Operculum colorless, translucent. Animal not observed. Habitat: Shallow subtidal, among algae. Distribution: Cape Verde Islands. Remarks: This species is closely allied to the previous one, but has a distinctive, Coriandria-like appearance with its small size, glossy, pupoid shell and distinct band- ing pattern. It is also distinguished from all other species treated here by the colorless, translucent operculum. The correct placement in Barleeia is ascertained by the char- acters of the protoconch, operculum and radula. This species is deilicated to Maria Josefa "C.hefi Al- varez Aza, Dr. Rolan s wife and a companion of the Cape Verde expeditions. Barleeia tomensis Gofas, new species (Figures 7-8, 39-40) Type material: Holotype (MNHN) and 24 paratvpes (12 MNHN, 12 MNCN cat. 15.05/20531) collected alive from the type locality. Type locality: Praia das Conchas, Sao Tome (00°24.7'N, 06°38.0"E), at lou tide. Material examined: Sao Tome: Esprainha, 50* + 58 specs., Gofas and Fernandes leg. 11.1985 (MNHN); Praia das Conchas, the type material, Gofas and Fernandes leg. 11.1985; Praia das Conchas, 13 sh., Rolan leg. (CER); Mutamba, 28 specs., Fernandes leg. 12.198b (MNHN); Mutamba, 53 sh., Rolan leg. (C^ER); Lagoa Azul, 5 sh., Rolan leg. (CER); Praia Morro Peixe, 2 specs., Gofas leg. 11.1983 (MNHN); Santa Ana, 2 m, 11 specs., Rolan leg. (CER). Description: Shell conical, solid, atlults 1.8 x 1.1 to 2.4 X 1.3 mm (holotvpe 2. 1 x 1.2 mm). Protoconch high dome-shaped (350 fx in diameter) of l'/2 to 1% whorls, with very minute spiral rows of pits. Teleoconch with 3'/4 to 3'/2 whorls. Spire whorls moderately convex, body whorl rounded and imperlorate. Outer lip slightly pro- socline, beveled inside to a thin edge; parietal callus rather thin. Color of shell reddish brown, paler towards the parietal insertion of the body whorl; typically with two periumbilical spiral bauds on body whorl. Opercu- lum dark crimson. Head-loot w ith superficial black pat- tern covering the snout, fading behind the head; on oper- cular lobes and anteriorly on the sides of the epipodial folds; whitish/yellow axial bar on tentacles, whitish/yel- low graiuilar masses behind each eye, on the tip of the snout, on the sole oi the foot and the propodinm, and on the opercular lobes anteriorly to the black markings. tiahltal: Intertidal to shallow subtidal on rock\ shores witli clear water; in algal growth oi coralline and en- crusting red algae. Distribution: Only known from the Islands of Sao Tome and Principe. Remarks: This species was cited by Tonilin and Shac- klefortl (1914) and Fernandes and Rolan (1993) as Bar- leeia rubra The spotted sole of the foot distinguishes it from B. gougeti and B. unifaseiata, which always have a colorless sole. The high spire recalls that of B. verdensis n. sp., but B. tomensis has a constant, characteristic dou- ble pale band around the umbilicus, and does not have the aperture beveled outside. Barleeia taeniolata Gofas, new species (Figures 9, 41-42) Type material: Holotype and 5 paratypes (MNCN cat. 15.05/20532), empty shells from the type locality. 5 para- types (sh.) from Lagoa Azul (MNHN) Type locality: Santa Ana, Sao Tome (00°15.5'N, 06°45"W), 2 m. Material examined: Sao Tome: Santa Ana, 2 m, 6 sh., Rolan leg. (holotype and paratypes MNCN); Baia de Ana Chaves, 2 m, 5 sh., Rolan leg. (CER); Praia das Conchas, 5 m, 1 sh., Rolin leg. (MNHN); Praia das Conchas, 2 sh., Gofas leg. (MNHN); Lagoa Azul, 4 m, 14 sh., Rolan leg. (paratypes MNHN; CER); Mutamba, 4 m, 21 sh., Rolh leg. (C;ER). Principe: Santo Antonio, 6-10 m, 1 sh., Rolan leg. (CER). Description: Shell conical, rather translucent, holotype 1.6 X 1.05 mm. Protoconch high dome-shaped (300 /x in diameter) of I'A whorls, with very minute spiral rows of pits. Teleoconch with 3 whorls. Spire whorls convex, body whorl rounded and imperforate. Outer lip slightK prosocline, thin; parietal callus rather thin. Color of shell of a pale horny color, with two brown bands on the spire whorls and two more on the bocK whorl, one continuing the suture and one [jeriumbilical; the two central bands on the body whorl commonly fused to form one broad band. Operculum and animal unknown. Habitat: I'nknown, [)resuinabl\ sublittoral in .shallow water. Distribution: Only known from the islamls of Sao Tome and Principe. Remarks: 'ilie pitted protoconch of this species is char- acteristic ot Barleeia although the shell resembles a Cor- iandria at first glance. It is easily separated irom jinenile B. tomensis by the shorter spire, more convex whorls, and different color pattern. Barleeia taeniolata always has a pale area separating the suprasutural and subsuturai browti bands on the whorls, whereas B tomensis only has periumbilical pale bands. Barleeia pervulgata Gofas, new species (Figures 10-15, 47-52, 71-72) Type material: ilolotspe (MNHN) and over 2000 para- types (500, ot which 100 are juveniles, in each of the S. Gofas, 1995 Page 21 Figures 10-12. Barlecia pervtilgata Gofas, new species. 10. Holotype from Chapeu Armado (Angola), length 2.2 mm. 11. Living animal from Chapeu Armado (Angola), length of shell 2,7 mm. 12. Egg capsule containing a larva ready to hatch, from Chapeu Armado (Angola). Scale bar 1 mm. Figures 13-15. Barleeia cf. pervulgata Gofas, new species. 13. Specimen of a "pale" morph from Santa Maria (Angola), length 2.1 mm. 14. Specimen of a subtidal morph from Santa Maria (Angola), length 2.4 mm. 15. Living animal of a subtidal morph from Porto Alexandre (Angola), length of shell 1.7 mm. Page 22 THE NAUTILUS, Vol. 109, No. 1 following: AMS cat. C.305091, MNCN cat. 15.05/20533, MNHN, UAN) collected alive from the type locality. Type locality: Chapeu Armado, province of Namibe, Angola {14°27'S, 12°21'E), intertidal. Material examined: (collected by the author; all MNHN e.xcept paratvpes specified above) Angola: (typical form) Caotinha, 2800+ specs. (1.8 x 1.1 to 2.4 x 1.4 mm); Limagens, 800 specs., 12.1981 (1.7 x 1.0 to 2.2 x 1.3 mm); Santa Maria, intertidal, 1000 specs., 12.1981 (1.8 X 1.1 to 2.2 X 1.3 mm, man\ pale specs); Santa Maria, dredged 8-10 m, 200 sh.; Lucira (Bissonga), 37 specs. (1.7 X 1.0 to 2.1 X 1.3 mm); Baia das Pipas, 300* specs., 2.1983 ( 2.4 X 1.5 to 2.9 x 1.7 mm); Sao Nicolau, 22* + 100* specs., 2.1983 (2.1 x 1.3 to 2.9 x 1.7 mm); Chapeu Armado, the type material (1.9 x 1.2 to 2.7 x 1.6 mm); Praia das Conchas, 200 specs. (2.2 x 1.4, to 2.8 x 1.7 mm); Praia Amelia, 400 + 18* specs., 2.1983 (1.6 X 1,0 to 2.5 X 1.4 mm, some white specs.), {pale, banded form) Caotinha, 2 specs. + 8 sh.; Limagens, 130 specs., 12.1981 (1.6 x 1.1 to 2.3 x 1.4 mm); Santa Maria, 0-2 m, 1 sh.; Sta Maria, 8-10 m, 12.1981 on calcareous algae: 50 specs. + 8 sh. (2.4 x 1.4 to 1.8 x 1.2 mm); Lucira (Bissonga), 2 specs.; Sao Nicolau, 7* + 25 specs. (2.1 X 1.2 to 2.2 X 13 mm); Chapeu Armado, 9 specs. + 24 sh. (2.0 X 1.3 to 2.7 x 1.6 mm); Praia Amelia, 19 specs. + 1 sh.; Baia dos Tigres, in mussel bed, 12*+ 3000 specs., 8.1985. (more delicate form from deeper water): Santa Maria, 8-10 m, 12.1981 on calcareous algae, 600 specs.; Baia da Lucira (Osar), 10 m, 10 sh.; Santa Marta, 40 m in shell sand, 100 sh.; Praia Amelia, 40-60 m, 25 sh.; off Porto Alexandre, 5* + 2 specs. (1.7 x 1.1 to 2.2 x 1.3 mm). Description: Shell conical, very solid, adults 1.7 x 1.0 to 2.9 X 1,7 mm (holotype 2.2 x 1.3 mm). Protoconch dome-shaped (350 n in diameter) of I'/s to 1% whorls, with very minute spiral rows of pits, sometimes also with ill-defined spiral sculpture. Teleoconch with 3'/4 to SVz whorls. Spire whorls rather flat, body whorl rounded and hardly unibilicate. Outer lip slightK prosocline, beveled inside to a thin edge; parietal callus tiiickened in adults, tending to form a ridge. Color of shell reddish brown, paler towards the parietal in.sertion of the body whorl; plain or with whitish spiral bands, subsutural, median on body whorl and/or periumbilical; rarely entirely oi a pale, wax-like color. Operculum dark crimson. Head- foot with superficial black pattern on upper part of head generally forming a V-shaped |)atlern pointing forwards, then extending over the snout to the sides and axiulK; two small black spots on the sides of the propodium, sometimes also a pair of larger, blurred, black to grayish spots on the axial part of the propodium; opercular lobes black. Yellow axial bar on tentacles; yellow granular masses behind each eye, on each lobe of the tip of the snout and on opercular lobes anteriorly to the black areas; sole of foot circled by a line of yellow granules. Habitat: Intertidal to shallow subtidal on rocky shores with clear water; in algal mats, mainly oi coralline algae. Distribution: Known only from the provinces of Ben- guela and Namibe, Southern Angola. Remarks: This is by far the commonest Barleeia of Southern Angola. It somewhat resembles the European species B iinifasciata but is readily diagnosed by the yellow patterns on the foot, absent in the former. The egg capsule (Fig. 12) has been observed in Chapeu Armado, and is very similar to that of B. unifasciata (Lebour, 1934; Southgate, 1982). It is spherical, contains a single developing larva, and is attached to the algal mat and embedded sand grains. The above description (and Figs 10-11) is based on the populations commonly found in the intertidal zone from Caotinha to Praia Amelia. Besides this, there are two forms which may be distinguished morphologically and are sympatric in most localities: (1) One form has a paler, commonly banded shell, with slightly more convex whorls and more distinct um- bilical chink (Fig. 13). Such specimens also commonK have a taller protoconch, with more conspicuous spiral pattern. They are found on the shore at slightly lower levels than typical B. pervtilgala, which is restricted to the shore. These forms look quite different at first glance, but many individuals cannot be conviiicingK separated within microsympatric populations; the sample from Limagens (intertidal) contains many banded forms of ambiguous morphology (e.g. with banded pattern but not with convex whorls). Also, the color pattern of the animal, usualK a useful character at species level, varies within the same limits in both forms. (2) A second form with a more delicate, taller, plain brown shell, occurring always subtidally on soft bottoms with calcareous algal concretions (Figs 14-15). This form never occurs microsympatrically with typical B. per- vulgata, but is found in Santa Maria together with the above "paler" form in 5-10 m, where both lorms cannot be convincingK separated. The color pattern of the an- imal is not very different from that of palest intertidal morphs. More study (e.g. using allozyme electrophoresis) may be necessary to determine whether the differences are merely ecophenotypic variation, or if there are more than one species with overlapping variation and/or limited introgression. Barleeia ciugiilnidcs CJolas, new species (Figures 16-17, 43-44) Type material: Holotype (MNHN) and M paratypes (16 of which 4 juv., in each of the following: AMS cat. C.305092, MNCN cat. 15.05/20534, MNHN, UAN)col- leited alive Irom the t\pf locality. Typ<' locality: Praia .Vmelia, province of Namibe, .Vngola (r5°13'S, 12°07'E), intertidal. Material examined: (collected by the author; all MNHN except paratypes specified above) Angola: Baia de Santa Maria 8-10 m, 5 sh.; Santa Marta, 40 m, 2 sh.; Baia das Pipas, 53* specs., 2,1983 (2.2 x 1.2 to 2.4 x 1.4 mm); S. Gofas, 1995 Page 23 Figures 16-17. Barlecia cinguluidc.s Gofas, new species. 16. Holotype from Praia Amelia (Angola), length 2.4 mm. 17. Living specimen from Praia Amelia (Angola), length of shell 2.4 mm. Figure 18. Barleeia picta Gofas, new species, holotype from Santa Maria (Angola), and operculum seen froiii outer side; length of shell 1.8 mm. Praia Amelia, intertidal, the type material (adults 2 2 x 1.3 to 2.5 X 1,4 mm) and 50 specs, (jiiv.). Description: Shell high conical, quite solid, translucent, adults 2.2 X 1.2 to 2.5 x 1.4 mm (holotype 2.4 x 1.3 mm). Protoconch high dome-shaped (350 n in diameter) of l'/2 to 1% whorls, with spiral rows of relatively coarse pits, some of them irregularly fused along one row. Te- leoconch with 3'/4 to 3'/2 whorls. Spire whorls rather flat, body whorl with a faint but sharp keel continuing the suture, imperforate. Outer lip nearly orthocline except for the parietal insertion where it is pro.socline, beveled inside to a thin edge; parietal callus moderately thick- ened. Color of teleoconch pale buff, with narrow brown lines, one subsutural, one suprasutural on the spire and running just above the keel on body whorl, and one periumbilical. Operculum pale brown, external surface shagreened with hardly visible growth lines. Head-foot with superficial black pattern forming two patches an- teriorly on each side of propodium, two longitudinal streaks on the snout and patches on opercular lobes. A solid yellow bar on tentacles, yellow granular masses behind eyes, small yellow spots on each lobe of the snout, and on the sides of propodium; sole colorless. Habitat: Intertidal on rock\ platforms, among rocks cov- ered with coralline algae and lying on coarse, clean sand. Distribution: Known only from the provinces of Ben- guela and Namibe, Southern Angola. Remarks: This species recalls the European rissoid Cin- giila cingilliis (Montagu, 1803) in shape, color pattern, and even the intertidal habitat in sand under stones. It is readily distinguished from other Barleeidae by its color pattern; banded forms of 6. pcrvitlgata never have such narrow lines and lack the peripheral keel. The most close- ly related species is B. picta n. sp., similar in shell shape and in having coarse punctures on the protoconch, but colored with flames instead of spiral lines. Page 24 THE NAUTILUS, Vol. 109, No. 1 Barleeia picta Gofas, new species (Figures 18, 45-46) Type material: Holotvpe (MNHN) and 37 paratypes (9 AMS cat. C.305093,'9 MNCN cat. 15.05/20535, 10 MNHN, 9 LI AN) collected alive from the type locality. Type locality: Baia de Santa Maria, province of Ben- guela, Angola (13°35'S, 12°33'E), on calcareous algal bot- tom in 8-10 m. Material examined: (collected by the author; all MNHN except paratypes specified above) Angola: off Ambrizete, 45 m, 7 sh. (2.2 x 1.4 to 2.6 x 1.6 mm, subfossil?); off Ilha de Luanda, 120 m: 1 sh (subfossil); Santa Maria, the type material (1.8 x 1.1 to 2.3 x 1.3 mm); Lucira (Santa Marta),12sh. (1.5 x 0.9tol.7 x 1.0 mm); Lucira (Caesar), 1 sh.; Sao Nicolau, 15 sh (1.8 x 1,0 to 1.9 x 1.1 mm); Ponta Albina/Baia dos Tigres, 40 m, 12 sh. Description: Shell conical, translucent, adults 1.5 x 0.9 to 2.3 X 1.3 mm (holotype 1.8 x l.Q mm). Protoconch dome-shaped (300 n in diameter) of l'/2 to 114 whorls, with ill-defined, Dat spirals and covered with spiral rows of rather coarse, pits. Teleoconch with 3'/4 to 4 whorls. Spire whorls flat, body whorl faintly angulated along a line continuing the suture, non-umbilicate. Outer lip or- thocline, with very thin edge, thickened in the adult at some distance from its edge. Color of shell very pale buft to pinkish on apical whorls; two last whorls with a series of very faint subsutural flames, a colorless zone around the periphery of the body whorl; another series ot taint flames on periumbilical area, matching the subsutural ones or merged into a continuous darker periumbilical band. Operculum dark crimson, along the ridge and edges, pale yellowish on distal surface; external surface shagreened with hardly distinct growth lines. Patterns of animal not observed in life, seen on rehydrated spec- imens to have black markings on opercular lobes and on the snout. Habitat: On bottoms of calcareous red algae ("maerl") in a few meters depth and clear water. Distribution: Known only from Angola; live-taken onl\' in the [irovince of Benguela, Southern Angola. Remarks: This species shares with B. cinguloidcs the sculpture of very coarse pits in the protoconch. Shells found on the continental shelf of Northern Angola are similar, but larger. They are presumably derived from Pleistocene deposits which occur in 50-120 m; the ma- terial in such ,sam|)lcs contains many littoral species, in- cluding some now restricted to the south Genus Pseudodiala Ponder, 1967 Type species: Diala acuta (Carpenter, 1864, by original designation. Shell with smooth, rather thin teleoconch, very narrov\ ly umbilicate, generalK whitish; spire whorls rather flat, bodv whorl with a faint peripheral keel. Aperture ovate. with a thin, opisthocline outer lip. Operculum as in Bar- leeia . Radula (Ponder, 1983 and herein. Fig. 73-74): central tooth with a large median cusp with blunt tip, and 2-3 small lateral denticles; one sharp basal denticle on each side, separated by a broad U-shaped projection. Lateral and marginal teeth as in Barleeia. Ponder (1983) synonymized Pseudodiala with Bar- leeia, but P. niso n. sp. shares with the type species of Pseudodiala, P. acuta, the opisthocline aperture (see Fig. 19) and the narrow central cusp of the radula. The West African species assigned to Pseudodiala also stand apart from Barleeia by their habitat on soft bottoms of the shelf rather than in intertidal or shallow subtidal hard bottom biota. These are retained as a separate group on these grounds. Pseudodiala niso Gofas, new species (Figures 19, 53-54, 73-74) Type material: Holotvpe collected alive (MNHN) and 55 paratvpes (15 AMS cat. C.305094, 20 MNCN cat. 15.05/20536, 20 MNHN), empty shells from the type locality. Type locality: Senegal, off Goree, 50 m (14°32'N, 17°25.5'W). Material examined: (all MNHN except paratypes speci- fied above) Senegal: off Goree, 50 m, the holotype and 55 paratypes (sh.), Marche-Marchad leg.; Cap de Naze, 25 m, from fish stomach, 3 specs. (3.2 x 1.8 to 3.5 x 2.0 mm), Marche-Marchad leg.; Off Saloum, 50 m, 28 sh., Marche-Marchad leg. ;'"Petite cote (i.e. coast be- tween Dakar and the Saloum estuar\ ), 32 m, 1 spec, Leung-Tack leg. Description: Shell conical, translucent, adults 2.7 x 1.5 to 3.5 X 2.0 mm (holotype 2.7 x 1.7 mm). Protoconch stub-shaped with rounded top (400 ^ in diameter) of l'/2 to 1% whorls, with imperforate surface, ill-defined, flat spirals and a very definite suprasutural keel. Teleoconch with 3'/2 to 4'/2 whorls. Spire whorls hardly convex, body whorl carinate along a line continuing the suture, dis- tinctk umbilicate Outer lip strongK opisthocline, bev- eled to a (juite thin edge, parietal callus separating from the body whorl in adults. Color of shell uniformly whitish to gravish. Operculum dark crimson, w ith a faint exter- nal ridge along the cohunellar siile. .Vniinal not observed. Habitat: On the continental shelf, 30-100 m Distribution: Known only from Senegal. Remarks: The imperforate protoconch of this species is at\pical in the Barleeidae, but this character state is bridged to the more usual pittetl protoconch by the tiny punctures of the next species P. aequiiinctialis n. sp. The iaint suprasutural keel in the protoconch ol P. niso is liiiinologous to the stronger suprasutural spiral ridge in /' acquinoctialis. and the overall siniilarit\ in proto- conchs indicates a close relationship between these spe- S. Gofas, 1995 Page 25 Figure 19. Pseudududa luso Ciofas, new spccii'S. li(ilot>pe from off Goree (50 m). Senegal, leiigtli 2 7 iiiiii. Figure 20. PscudodUila aequinoctialis Gofas, new species, holotspe from off Equatorial Cknnea (1.50 ni). lengtli 1.6 nun Figure 21. Pseudodiala corollaria Gofas, new species, liolot> pe from Santa Ana, Sao Tome, lengtfi 1.7 mm. Figure 22. Tropidormuia taphrudcs Tomlin & Sfiackleford, 1915, specimen from Praia das Conctias, Sao Tome, length 1.7 mm Figure 23. Twpidorissoia secttnda Rolan & Templado, 199-4, paratype (MNHN) from Praia Emilia, Sao Tome, length 1.1 mm. Page 26 THE NAUTILUS, Vol. 109, No. 1 cies. Tlie name is allusive to the resemblance with the eulimid genus Niso. Pseudodiala acquiiwctialis Gofas, new species (Figures 20, 55-56) Type material: The holotype (MNHN). Type locality: Equatorial Guinea (0r40'N, 09''25'W), 150 m. Material examined: Equatorial Guinea: the holotype. Principe: Santo .Antonio, 6-10 m, 21 sh., Rolan leg. (CIER); Baia das Agulhas, 8 m, 21 sh., Rolan leg. (CER). Description: Shell conical, translucent, adults 1.6 x 1.0 to 2.4 X l.,3 mm (holotype 1.6 x 0.9 mm). Protoconch globular with rounded top (300 fi in diameter) of 1 '/2 whorl, with broad spiral ridges, the lowermost of which forms a suprasutural keel, and tiny punctures evenly distributed without lineations. Teleoconch with 3'/2 whorls, externally smooth, sometimes with irregular in- ternal punctures seen by transparency. Early spire whorls somewhat convex, body w horl carinate along a line con- tinuing the suture, distinctly umbilicate. Outer lip strong- ly opisthocline, thin. Color of shell uniformly whitish to grayish. Animal and opercuhun not observed. Habitat: Unknown, holotype collected as an empty shell on the continental shelf (150 m depth), from a geotechni- cal core for petroleum exploration, additional material dredged in (i-10 m on soft bottoms around the island of Principe. Distribution: Known from Equatorial Guinea and the island ot Principe. Remarks: This species is allied to P. niso. with which it shares the general shape, opisthocline lip (broken on all specimens but inferred from the growth lines), the keeled and umbilicate body whorl. It is readily distinguished from P. niso by its protoconch with distinct spirals and tiny perforations. It is distinguished from the next species P. corollaria n. sp. by lacking the brown color on the parietal side of the aperture, by its larger size, higher spire, flatter whorls and by the profile of its last whorl, which is slightly angulose (more conspicuously in im- mature specimens). The holotype, collected in a box core, is slightly im- mature and possibly derived from a Pleistocene deposit, but is the best preserved specimen. On some specimens from Principe, some tiny and evenly distributed jjunc- tures can be seen by transparency inside the spire w horls This feature may be restricted to some specimens and could not be .seen on the holotype. Pseudodiala corollaria (iofas, new species (Figures 21, 57-.58) Type material: Holotype (MNCN cat. 15.05/20.536) and 7 paratypes collected alive from the type locality. 3 para- types (MNHN) collected alive from Praia Mutamba. Type locality: Santa Ana, SSo Tome (00°15.5'N, 06°45'W), 2 m. Material examined: (all collected by E. Rolan) Sao Tome: Santa Ana, 2 m, 8 specs., (holotype and paratypes MNCN)and 3 sh.(CER); Praia Mutamba, 4 m, 3 specs., (paratypes MNHN); Lagoa Azul, 4 m, 5 sh. (CER); Praia das Conchas, 5 m, 6 sh. (C>ER). Description: Shell conical, translucent, adults 1.4 x 0.8 to 1.8 X 1.0 mm (holotype 1.7 x 1.0 mm). Protoconch globular with rounded top (300 n in diameter) of l'/2 whorl, with broad spiral ridges, and tin\ irregular pits, very loosely arranged along spiral rows. Teleoconch with 3 to 3'/2 whorls, externally smooth, sometimes with ir- regular internal punctures seen by transparency. Spire whorls quite convex, body whorl rounded, distinctK um- bilicate. Outer lip strongly opisthocline, thin. C^olor of shell whitish to grayish, translucent, with the parietal edge of the aperture strongly tinged with brown. Pro- toconch generally more opaque, tinged with \ellowish or reddish. Operculum dark crimson. Animal not ob- served; remains of a black mantle seen attached to the interior of the shell. Habitat: On soft bottoms of muddy sand in shallow sub- tidal sites. Distribution: Known from the Island of Sao Tome. Remarks: This species differs from the closely related P. aequinoctialis by not being carinate at any stage, by its smaller size and more convex whorls, b\ the more distinctly pitted protoconch, and by the dark columellar edge of the aperture. Genus Tropidorissoia Tomlin and Shackleford, 1915 Type species: Tropidorissoia taphrodes Tomlin and Shackleford, 1915, by monotypy. Shell with stout, solid teleoconch and strong spiral keels, very narrowly umbilicate, generally tinged w ith brown. Aperture ovate, slightly prosocline to orthocline, with smooth imier lip, thickened at some distance inside and then beveled to a thin edge. Protoconch with spiral series of pits, with or without superimposed spiral cords. Head- foot pigmented with black and/or yellow. Operculum as in Rarleeia excejit lor pale brown color. Radula (Fig. 75-76) as in Barlecia. All the species from St. Helena (Rissoa acqua, R. per- fccta, R. plotia, R. varicifera, R. vaga, R. ivallichi, R. simulaiis. all of Smith, 1890) classified or tentatively classilied in Tropidorissoia by Tomlin and Shackleford (1915) were referred or tentatively referred by Ponder (1985) to his new ris.soid subgenus Lirocingula. on the basis of conchological siinilaril\ w ith the South African t\[)e species /.. winslowae (Bartsch, 1928). Data on the soft parts, radulae and opercula are at present wanting for a definitive placement. S. Gofas, 1995 Page 27 Tropidorissoia taphrodes Tonilin ami Sliacklt'tord, 1915 (Figures 22, 59-60, 75-76) Trupidurissiiiu laplixnlcs roiiillii and Sliacklt'tord, 191.5 pi. 5, tie. 5 Type material: llolotype (live collected) in British Mu- seum (Natural History); 15 paratypes in National Mu- seum of Wales, Cardiff, cat. 1955.158.1121. Type localily:Isiand of Sao Tome. Material examined: Sao Tome: Esprainha, 19* -t- '30 specs., Gofas and Fernandes leg. 11.1985; Praia Mutam- ba, 9 specs., Fernandes leg. 12.1986; Praia Mutamba, 6 sfi., Rolan leg. (CER); Lagoa Azul, 5 m, 5 sh., Rolan leg. (CER); Praia das Conchas (Guadalupe), 19* + 40 specs., Gofas & Fernandes leg. 11.1985; Praia das Conchas, 11 sh., Rolan leg. (CER); Baia de Ana Chaves, 4 sh., Rolan leg. (CER); Santa Ana, 3 specs. Rolan leg. (CER). Prin- cipe: Santo Antonio, 6-10 m, 41 specs. Rolan leg. (C!ER); Baia das Agulhas, 8 m, 19 specs,, Rolan leg. (C]ER). Description: Shell conical, solid, adults 0.8 x 0.5 to 1.2 X 0.7 mm. Protoconch dome-shaped (350 p in diameter) of l'/2 whorl, with strong spiral cords and superimposed minute spiral rows of pits. Teleoconch of 2 to 2'/i; whorls, with two strong, rounded keels on the spire whorls and with a third similar keel on the body whorl; a spiral thickening around the small umbilical chink. Outer lip slightly prosocline, thickened deep inside, then beveled to a thin edge. Parietal callus moderately developed, detached from the umbilical chink. Color of shell reddish brown, sometimes pale yellowish. Operculum yellov\, translucent, with an internal peg. Head-foot with su- perficial gra\ pattern on the snout and opercular lobes. Opaque vellowish white flecks a.xially on tentacles, tri- angular opaque yellowish-white granular masses behind each eye. Reddish buccal mass clearly seen by transpar- ency. Foot white, with tiny opaque white flecks on the sole; metapodium not conspicuously cleft as in Barlccio. Habitat: Intertidal to shallow subtidal on rocky shores with clear water; in algal mat, mainly of coralline algae. Distribution: Known only from the islands of Sao Tome and Principe Remarks: Fernandes ami Rolan (1993) noted a differ- ence in the populations from Principe, these having fre- quently a pale shell whereas all specimens from Sao Tome are uniformly reddish-brown. In the material ex- amined, 36 out of 41 specimens from Santo Antonio and all 19 specimens from Baia das .Agulhas are pale. The protoconch has a peculiar sculpture, cjuite differ- ent from that of most Barleeia, but similar to that of " Barleeia" congenita Smith, 1890 from St. Helena Island (Fig. 63-64). Tropidorissoia secunda Rolan and Templado, 1994 (Figures 23, 61-62) Tritpulfjrissoiu sccuruhi Holan ami 'rcinplado, 1994: 2.37-242, liU 1-1 Type material: llolotype (MNCN cat. 15.05/6975); Paratypes (3 MNHN, 3 British Museum (Natural His- tory), 3 American Museum of Natural History, 10 CER, 10 private collection of F". Fernandes, Luanda). Type locality: Praia Emilia, Sao Tome (0O°21.7'N- 06°43.5'E), Material examined: (collected by E, Rolan) Sao Tome: The MNHN paratypes; Praia das Conchas, 6 sh. (CER); Lagoa Azul, 4 m, 25 sh., (CER); Baia de Ana Chaves, 1 sh. (C;ER). Principe: Santo Antonio, 6-10 m: 17 sh. (CER); Baia das Agulhas, 8 m, 25 sh. (CER). Description: Shell conical, solid, adults 0.8 x 0.5 to 1.2 X 0 7 mm. Protoconch dome-shaped (220 yu in diameter) of 1 '/2 whorl, with minute spiral rows of pits. Teleoconch of 2 to 2'/2 w horls, with a strong median keel on the spire whorls and with a suprasutural cord which extends on the bnd\ whorl to form another keel. Outer lip slightly prosocline, beveled inside to a thin edge. Parietal callus moderately developed, detached to form an umbilical chink. Color of shell white to pinkish. Operculum yellow, translucent. Head-foot with superficial plain black pat- tern on the snout and opercular lobes. Opaque white axial bar on tentacles, large, triangular opaque white granular masses behind each eye. Pink buccal mass vis- ible by transparency. Habitat: Subtidal in a few meters depth. Distribution: Only know n from the islands of Sao Tome and Principe, Genus Liroharleeia Ponder, 1985 Type species: Alvania nigrcscens Bartsch & Rehder, 1939 (Alvania galapagensis Bartsch, 1911 sensu Ponder, 1983), pending decision of ICZN (see Hertz, 1994 for a discussion of the type species). Shell with elongate, solid teleoconch and spiral sculpture in all species considered, non-umbilicate or very narrow- ly umbilicate, generally tinged with brown. Protoconch with minute pits loosely arranged in a spiral direction in the species considered herein; with strong spiral ridges and no pits in the type species. Outer lip opisthocline like in most Rissoina species. Head-foot pigmented with black and/or yellow. Operculum as in Barleeia. Radula (Ponder, 1983 and herein. Fig. 77-78): central tooth with a large rectangular median cusp, one large triangular denticle on each side of it, and sometimes one or two very small denticles flanking these on the sides of the tooth; one sharp basal denticle on each side, separated by a broad, prominent lamella. Lateral teeth (one pair) with a rather broad base, terminating with a large, rather trapezoidal cusp flanked by slightly smaller cusps to ei- ther side; with a U-shaped projection beneath those. Mar- Page 28 THE NAUTILUS, Vol. 109, No. 1 ginal teeth (two pairs) narrow, hook-shaped, with uni- pectiiiate small cusps towarils the distal end. I have included the three species described here in Liroharleeia despite important differences in both pro- toconch and teleoconch scupture. They are congeneric with L. albolirata (Carpenter, 1864) and L. kelscyi (Dall & Bartsch, 1902), two West American species that Ponder (1983) included in Lirubarleeia notwithstanding the same differences. The radula of the two species described here- in agrees fairly well with those illustrated by Ponder (1983), although lateral outermost denticles are wanting in the central tooth of L. siiblaevis n. sp. The specimen figured by Ponder (1983) to illustrate the designated type species Alvania galapagensis Bartsch, 1911 has been shown by Hertz (1994) to belong to Alvania nigreacens Bartsch & Rehder, 1939, and this case of misidentification should be settled by a decision of ICZN. The actual A. galapagensis is a rare species from deeper water and ma\ be a pickworthiid. The type species of Fictonoha Ponder, 1983 also re- sembles the species dealt with here, but differs in having a very different central tooth on the radula. Liroharleeia data Gofas, new species (Figures 24, 65-66) Type material: Holotvpe (MNHN) and 200 paratypes (50 AMS cat. C.305095, 50 MNCN cat. 15.05/20538, 50 VINHN, 50 UAN) collected alive from the type locality. Type locality: Baia de Santa Maria, province of Ben- guela, Angola (13°35'S, 12°33'E), on calcareous algal bot- tom in 8-10 m. IVIaterial examined: (collected by the author; all MNHN except paratypes specified above) Angola: Caotinha, 1 sh.; Baia de Santa Maria, the type material; Baia da Lucira (Cesar) on calcareous algal bottom 10 m, 1 spec, and 12 sh.; Santa Marta, dredged 40 m on shell sand, 25 sh.; Chapeu Armado, 1 sh.; Praia Amelia, 40-60 m, 3 sh. Description: Shell elongate, solid, adults 3.2 x 1.6 to 6.2 X 2.6 nun (holotype 5.1 x 2.3 mm). Protoconcli dome- shaped (500 n in diameter) with 1 to l'/-i whorl, a[)par- ently smooth at low magnification, with very minute, irregular pits, arranged in spiral bands so as to suggest a faint spiral sculpture. Teleoconch with 5'/4 to 6 whorls, with spiral sculpture of very uneejual, sharp cords (15- 20 on penultimate whorl). Early spire whorls rather flat, the later ones moderately convex, body whorl rounded, hardly umbilicate, with its latest part wider and salient from the profile. Outer lip opisthocline in adults, thick- ened at some distance in the last 'A whorl and beveled inside to the edge; parietal callus moderatcK developed, detached anteriorly in adults. Color of shell pale brown to whitish, with very faint, delicate axial flames starting from the suture; paler towards the termination of the body whorl; inside ol a()erture whitish to brown, often with two brown streaks on the lip. Operculum dark crim- son. Patterns of animal not observed in liie, seen on rehydrated specimens to have black markings on oper- cular lobes and on the snout. Habitat: On bottoms of calcareous red algae ("maerl") or coarse, clean shell gravel in a few meters depth in sheltered bays with transparent waters. Distribution: Only known from Angola. Remarks: There is some similarity with Rissoa glypta Smith, 1890 from St. Helena, which is smaller, has weak axial folds and somewhat carinate whorls. Liroharleeia data is the largest of the Angolan Liroharleeia, it is distinguished from both L. pupoides n. sp. and L. siib- laevis n. sp. by its more acute apical spire angle, and coarser sculpture; from L. siiblaevis also by the black markings on the animal and by the protruding lip of the aperture in the adults. Liroharleeia sublaevis Gofas, new species (Figures 25-26, 67-68, 77-78) Type material: Holotype (MNHN) and 480 paratypes (120 of which 20 juv. in each of the following: AMS cat. C.305091, MNCN cat. 15.05/20540, MNHN, UAN) from the type locality, live collected. Type locality: Praia das Conchas, province of Namibe, Angola (15°07.0"S, 12°06.7'E), in large tidal pools. IVIaterial examined: (collected by the author; all MNHN except paratypes specified above) Angola: Caotinha, 8 specs.; Limagens, 4 sh.; Baia de Santa Maria, 8-10 m, 500 sh.; Lucira (Cesar) 8-10 m, 150 sh. (2.6 x 1.4 to 4.3 X 2.2 mm); Lucira (Bissonga) intertidal 12* -I- 18 specs.; Santa Marta, 3 sh.; Sao Nicolau 14* specs. -I- 30 sh., 2.1983. Clhapeu Armado, 100 sjiecs. (many juv.); Praia das Conchas, the type material and 8* specs. (2.8 x 1.4 to 4.6 X 2.3 mm); Praia Amelia, 52 specs. (2.5 x 1.4 to 3.6 X 1.6 mm). Description: Shell elongate, very solid, adults 2.5 x 1.4 to 4.6 X 2.3 mm (holotype 3.0 x 1.5 mm). Protoconch dome-shaped (400^1 in diameter) of I'j whorls, appar- entlv smooth at low magnification, with extremely min- ute, irregular pits, loosely arranged along a spiral direc- tion. Teleoconch of 3V2 to 4'/2 whorls, with faint, flat spiral threads, the intervals of which show spiral series ot punc- tures (only visible under SEM). Whorls moderateK con- vex, body whorl rounded, slightK constricted, hardly umbilicate. Outer lip orthocline, thickened at some dis- tance in the last '4 w horl and beveled inside to the edge; parietal callus rather thin, (.letached anteriorly. C-olor of shell whitish or pale orange brown with subsutural area paler; occasionally w ith two blurred spiral orange brown bands (jn a whitish background. Operculum dark crim- son. Head-loot with \ellow axial bars on tentacles, yellow granular masses behind each eye, on the tip oi the snout, on each side of the anterior part of the propodium. Pink buccal mass clearK \ isible by transparency. Habitat: intertitlal, in pools or flats with rocks embedtled in clean, coarse sand and algal overgrowth. S. Gofas, 1995 Page 29 Figure 24. Lirobarleeia data Gofas, new species, holotype from Santa Maria (Angola), length 3.1 mm Figures 25-26. Lirobarleeia sublaevis Gofas, new species. 25. Holotype from Praia das Conchas (Angola), length 3.0 mm. 26. Living specimen from Caotinha (Angola), length 3.1 mm Distribution: Only known from Angola. Remarks: Lirobarleeia siihlacvis iTia> resemble some very smooth specimens in the population from Corimba, re- ferred to L. piipoides n. sp., but may be distinguished by its lack of black patterns on the head-foot, the absence of any pigmentation on its opercular lobes, and the reg- ularly fusiform profile of its shell, w Inch lacks the pro- truding body w horl. Lirobarleeia pupoides Gofas, new species (Figures 27-28, 69-70) Type material: Holotype (MNHN) and 200 paratypes (50 AMS cat. C.305096, 50 MNCN cat. 15.05/20539, 50 MNHN, 50 UAN) collected alive from the type locality. Type locality: Corimba, province of Luanda, Angola (8°50'S, 13°09'E), on shell gravel bar in 20 m depth. Material examined: (collected by the author; all MNHN except paratypes specified above) Angola: Corimba, the type material (2.2 x 1.1 to 3.8 x 1.5 mm); Corimba, Praia Etambar in shell sand: 34 shells (2.5 x 1.1 to 3.4 X 1.5). (Lirobarleeia cf. pupoides) Baia dos Tigres, 8* -I- 31 specs., 8.1985. (2.4 x 1.1 to 2.8 x 1.2 mm). Page 30 THE NAUTILUS, Vol. 109, No. 1 Figures 27-28. Lirobarleeia pupoides Gofas, new species. 27. Holotype from Oirimba (Angola), length 2.6 mm 28. Living specimen from C^oriniba (Angola), lengtfi 3.0 mm. Description: Shell pupoid, elongate, moderately solid, adults 2.2 X LI to 3.8 x L5 mm (holotype 2.6 x L2 mm). Protoconch dome-shaped (400 n in diameter) of I'/i whorl, apparently smooth at low magnification, with extremely minute, crowded, irregular pits, loosely ar- ranged along a spiral direction. Teleoconch of 3'/2 to AVi whorls, with flat spiral threads, the intervals of which show spiral series of punctures (only visible under SEM), and with very faint axial wrinkles. Whorls flat to mod- erately convex; body whorl rounded, hardly umbilicate. Outer lip orthocline, beveled inside to the edge which is quite rounded in adults; parietal callus moderately de- veloped, somewhat detached anteriorly and posteriorly. Color of shell orange brown to pale yellowish, occasion- ally with two blurred spiral bands inside the aperture. Operculum dark crimson. Head-foot with superficial blaik blotches on each side of the snout, occasionally confluent and extending over the head; black on oper- cular lobes and anteriorly on the sides of the propodium; yellow axial bar on tentacles, yellow granular masses behind each eye, on the tip of the snout, on the anterior part of the propodium, and on the opercular lobes an- teriorly to the black markings. Habitat: In coarse shell sand, shallow subtiilal. Distribution: Only known from Angola. Remarks: Very smooth specimens from C-'orimba may resemble the previous species, L. sublaevis in having a similar protoconch and teleoconch microsculpture. Li- robarleeia siihlacvis generally has a thicker and larger shell, a more fusiform profile without a protruding lip and lacks any black pattern on the head-foot and the opercular lobes. Old shells, possibly subfossil, collected on the beach at C^orimba, are larger (up to 4.3 x IS mm) and have a coarser spiral sculpture on the teleo- conch, approaching that of L. elata n. sp. They never- theless retain the pupoid outline that is typical of this species. In the extreme South of Angola, in Baia dos Tigres, there is a local population of Lirobarleeia resem- bling that of Clorimba both in shell morphology and color pattern of the head-foot, and also living in ca. 10 m depth on a coarse shell sand. It is not clear, from the material available, if it is the same species with a disjunct range, occurring where adequate substrate exists, or if it is a sister species derived independently from an ances- tral stock. DISCUSSION At the local level, sympatric species of Barleeidae may show differentiation in size, or in habitat. The NW Af- rican species Barleeia unifasciata and B. gougeti may be microsympatric and have distinct, although overlap- ping size ranges. The three Cape Verde Islands species described herein (B. verdensis n. sp., B. aemilii n. sp., B chefiae n. sp.) and the two Barleeia from Sao Tome (B. tomensis n. sp., B. taeniolata n. sp.) also exhibit size differentiation among microsympatric species. The svnipatric occurrence of several species (up to four in Baia de Santa Maria, Angola) is accompanied by a .segregation in habitat. Among the .Angolan species, Barleeia pervulgata n. sp. (the typical form) is found intertidally in algal tufts, B. cinguloides u. sp intertiilally among stones lying on coarse sand, while 6. pieta n. sp. occurs subtidally among algal concretions together with Barleeia cf. pervulgata and Lirobarleeia elata n. sp. The two Southern species of Lirobarleeia are also .segregated, L. sublaevis n. sp. occurring together with B. einguloides or in tidal pools, while L. elata is always subtidal. There is a preference for soft bottoms among Lirobarleeia and Fseudoitiala, and for hard bottoms among Barleeia. Thus lar, the coast of Western Ameriia was known as S. Gofas, 1995 Page 31 Figures 29-40. Protoconchs of Barleeia general views (scale bars 100 n) and enlarged portions of the surface (scale bars fO m), the latter all to scale, 29-30. Barleeia umjaseiata (Montagu, 1803), Ploumanac'h (France). 31-32. Barleeia gougeti (Michaud, 1830), Essaouira (Morocco). 33-34. Barleeia verdensis Gofas, new species, paratype, Salmanza (Sao Vicente, Cape Verde Is.). 35- 36. Barleeia aemilii Gofas, new species, Salmanza (Sao Vicente, Cape Verde Is.). 37-38. Barleeia chefiae Gofas, new species, paratype, Sai-Rei (Boavista, Cape Verde Is.). 39-40. Barleeia tomensis Gofas, new species, paratype, Praia das Conchas (Sao Tome). Page 32 THE NAUTILUS, Vol. 109, No. 1 Figures 41-52. Protoconchs of Barleeia: general views (scale bars 100 fi) and enlarged portions of the surface (scale bars 10 m). the latter all to scale. 41-42. Barleeia taeniolata Gofas, new species, paratype, Santa Ana (Sao Tome). 43-44. Barleeia cinguloides Gofas. new species, paratvpe. Praia Amelia (Angola). 4.5-46. Barleeia picta (lofas, new species, paratype, Santa Maria (Angola). 47-48. Barleeia pcrvnlgata Gofas, new species, (;:aotiiiha (Angola). 49-50. Barleeia pervulgata Gofas, new species, paratype, Chapeu Armado (Angola). 51-52. Barleeia cf. pervulgata Gofas, new species, Porto Alexandre (Angola). S. Gofas, 1995 Page 33 Figures 53-64. Protoconchs of Pseudodiaia, Tropidorissoia and "Barleeia". general views (scale bars 100 n) and enlarged portions of the surface (scale bars 10 n), the latter all to scale. 53-54. Pseudodiaia niso Gofas, new species, off Saloum (Senegal). 55-56. Pseudodiaia aequinoctialis Gofas, new species, holotype. Equatorial Guinea. 57-58. Pseudodiaia coroUaria Gofas, new species, paratype, Santa Ana (Sao Tome). 59-60. Tropidorissoia taphrodes Tomlin & Shackleford, 1915, Praia das Conchas (Sao Tome). 61-62. Tropidorissoia secunda Rolan & Templado, 1994, Praia das Conchas (Sao Tome). 63-64. "Barleeia" congenita Smith, 1890, St, Helena. Page 34 THE NAUTILUS, Vol. 109, No. 1 Figure§ 65-70. Protoconchs of Lirobarleeia general views (scale bars 100 /i) and enlarged portions of the surface (scale bars 10 ^l), tfie latter all to same scale. 65-66. Lirobarleeia elata Gofas, new species, paratype, Santa Maria (Angola), 67-68. L. suhlaevis Gofas, new species, paratype, Praia das Oinchas (Angola). 69-70. L. pupoides Gofas, new species, paratype, Corimba (Angola). the part of the world with the highest species richness in the family Barieeidae. Ponder (1983) recognized 13 species of Barleeia (including one Psetidodiala) distrib- uted between California and Panama, and 12 species of Lirobarleeia , distributed in the same mainland area but also in the Galapagos and Clarion islands. The description of species from West Africa brings attention to other centers where the family is well rep- resented (Fig. 79). The inner part of the Gulf of Guinea hosts si,\ species representing three genera. The absence of records from Annobon or Fernando Poo islands reflects the lack of data, rather than a real difference between islands. Southern Angola, with six species representing two genera, is another center of richness. The ne)rllu'rn limit of five species in Angola coincides with the shift from a rocky coast with clear, cool waters, to a prevalently sandy coastline with warmer, turbid waters north of the city of Benguela. Rocky shores with a well developed cover of small photophiious algae (including C'orallina spp. ) are well represented in the south, unlike the north where hard substrates are scarce and where high turbidity of waters would smother a dense algal cover. The sampling effort has been well balanced between l)()th areas, and the lack of records ot Barleeia in northern Angola is not a sampling artifact. This supports the view that the gaps between Angola and the Gulf of Guinea islands, and between the Gulf of Guinea and Senegal, are real. The distribution of Barleeia in West Africa may thus reflect the breakdown of a formerly more continuous range. To the south, the fauna of Namibia remains virtually unknown, lint some of the Angolan species, or related ones, may be present there; both Barleeia and Lirobar- leeia are present in the southernmost Angolan sample, less than 100 km from the Namibian border. Despite being a temperate area with rocky shores. South Africa has only two poorly know n species, Barleeia caffra (Sow- erby, 1897) and B. stnithi Bartsch, 1915, w hich are def- initely outside the West African radiation. The island of St. Helena may host a number of species of Barieeidae. However, the systematic position of most of the species described by Smith (1890) remains uncer- tain. Barleeia eongenita Smith, 1890 lacks spiral sculp- ture and superficially resembles true Barleeia, but its protoconch sculpture (Fig. 63-64) is similar to that of Tropidorissoia taphrodes. The species described by Sniilli as Rissoa and tentatively referred by Tomlin and Shac- kleford (1915) to Tropidorissoia may be either rissoids or barleeids (see under Tropidorissoia). Considering the Gulf of Guinea and Angola together, there are fewer species (12 species) than in West America but these are more diverse at the generic level: Tropi- S. Gofas, 1995 Page 35 ^^1 B- *^ ft««Bli^0L. livik ^ ^k^^^* ^^^^^^^^^ ^D t^^ ^i^i ^^ Figures 71-78. Kadulae, general views ot several complete rows (left) and enlarged views of central and lateral teeth (right; all scale bars 10 )i). 71-72. Barlccia pervulgata Gofas, new species, Chapeu Armado (Angola), 73-74. Pseudodtala niso Gofas, new species. Cap de Naze (Senegal). 75-76. Tropidorissoia taphrodes Tomlin & Shackleford, 1915. Praia das Conchas (Sao Tome). 77-78. Lirnbarlccia snblacvi.s Gofas, new species, Praia das Conchas (Angola). Page 36 THE NAUTILUS, Vol. 109, No. 1 Barleeia 1 .sp, \fi' Barlteui 'J ^-- 1 or 2 spp '.A^ Barleeia 3 spp. X Pseutioduila Isp Figure 79. Distrihiitidii ol the Barleeidae in the Eastern At- lantic: dashed contour, Barleeia, hatched, Tropidorissoia, and Liriiharlceia: stars, Pseiuhnhala dorissoia is present in addition to the three genera found in West America. Tluis, West Africa as a whole is the area of tfie world with the highest number of genera in the family. The representation in .Senegal (three species, two genera) and in the Cape Verde Islands (three species, all congeneric) is less spectacular Larval development without a planktonic stage has been observed only in B. unifasciata and in B. perviil- gata n. sp., but can be extrapolated to all the other species with similar protoconch characters. Despite this con- straint, the insular species are able to disperse between moderately distant islands. The three Barleeia species from the Cape Verde were collected on si.x islanils 10 to 100 km apart, and would i)rohal)ly have been foimd everywhere in the archipelago had all ten islands been sampled. The two species of Tropidorissoia and Barleeia tomenais described from Sao Tome were also foimd in Principe, at a distance of 150 km (Kernaiides and Kolan, 199.3) Conversely, there seems to be an allopatric dis- tribution for I'seudodiala acqttinuctialis u. sp., foimd in the inner part of the (Julf of Guinea, and P. corollaria found in Sao Tome. This implies that an efficient alter- native to dispersal during pelagic larval development is used by some barleeids. The most likely scheme is that the egg capsules of Barleeia, which are very light and attached to algae in shallow subtidal environments, are easily torn off the substrate and float away. ACKNOWLEDGMENTS The author thanks Jose Templado (MNCN) and Emilio Rolan (of Vigo, Spain) for providing the Cape Verde material, Anders Waren (Swedish Museum of Natural History, Stockholm) for preparing and scanning the rad- ulae, Philippe Bouchet (MNHN) and anonymous ref- erees for helpful conunents. The SEM micrographs of protoconchs were taken in liniversity of Malaga (Spain) by J. -J. Cuenca and G. Caballero, or in Centre Interu- niversitaire de Microscopic Electronique (Paris) by Mrs. D. Guillaumin. LITERATURE CITED Bellon-Hunibert, C. 1974. Les Molhisqnes testaces niarins du Maroc. Premier supplement, Travaux de Ihistitut Scien- tifique Cherifien, 37:l-l-t4. Borja, A 1986a. La alimentacion y distribucion del espacio en tres moluscos gasteropodos: Rissoa parva (da Costa), Barleeia unijaxiata (Montagu) > Bitliuni reticulatum (da Costa). C^aliiers de Biologie Marine 27:69-75. Borja, A. 1986b. Biologia > ecologia de tres moluscos gaster- opodos intermareales: Risnua parva. Barleeia unijaseiata y Bittium reliculatum. I Estructura y dinamica de las poblaciones. (^ahiers de Biologie Marine 27:491-.507. Borja, A 1987. Biologia y ecologia de tres moluscos gaster- opodos intermareales; Rissoa parva, Barleeia unifasciata y Bittium reticulatum. 2. (^recimiento ('ahiers de Biol- ogie Marine 28:351-360. Cosel, R. von, 1982. Ergebnissedeulscli-portuguesisclierSam- melreisen auf den Kap\erdischen In.seln (Repuhlica de Cabo Verde). Vorlaufige Liste der niarinen Mollusken. Courier Eorschung-lnstitut Senckenberg 52:15-25 Fernandes, F. and E. Rolan. 1993. Moluscos niarinos de Sao Tome y Principe: .■\ctualizaci6n bibliografica \ nuevas aportaciones. Iberus ll(l):31-47. Hertz, J. 1994. Review of the tvpe species ol Liroharlecia Ponder, 1983. The Veliger 37(1): 1 10-1 16 Lebour, M,\'. 1934. Ri.ssoid larvae as lood ol the \,iliacjoiinis. 6. Fiilly-grown, partly decollated shell 7. Dt-tail of another specimen, with the protDciinch present, shovvuig an extremely thin area (arrows) above the septum. 8-10. Tahcilia licinrichi. 8. Juvenile 9. Decollated and 10. non-decollated adult shells (arrow indicating extremely thin area). Scale bars = 2 mm. below the septum. Above the septum, the shell whorls, or what remained of them in partially decollated spec- imens, were always as thin as in conspecific juvenile specimens. These observations suggest that, in a first phase, these snails produce all the shell whorls, one after the other, of about equal thickness. Once the shell wall has been formed, there is no additional thickening in this initial phase of shell growth, which ends with the formation ot the characteristic body whorl, thought to be indicative of adult size. In a second phase, either the entire shell is thickened from within (Figure 3), or an apical septum is formed and the secondary thickening is restricted to the whorls below this septum (Figure 4). The latter cat- egory contains shells with relatively thin-walled apical whorls, prone to irregular breaking, usually resulting in conspicuously decollated shells (F"igure 4: slightly decol- lated). In Albinaria, septum formation is usually followed by decollation, as it is in A. rebeli Wagner, 1924. This species occurs abundantly in a small valley in Crete, where the snails can easily be observed, hanging on the rock-faces. Among lumdreds of full-grown, decollateil specimens, six adult shells with the apical whorls still present were collected. These were X-rayed. All of them contained an apical septum, above which the whorls are thinner than below it (Figure 5). No traces of shell wall resorption were seen. Septum formation may also be related to incidental damage of the apical whorls, as is most probably the case in A. spratti (Pfeiffer, 1846). Upon X-ray examination, an apical septum was observed in 6S of 208 shells of this species. Either decollation or fairly serious damage to the uppermost whorls was observed in these specimens. The remaining 140 specimens were undamaged and lacked a septum. Fully grown specimens of A. spratti may have a relatively thick shell wall from the body whorl up to the protoconch (Figure 3). In both A. rcbcli and A. spratti, septum formation might be hypothesized to be a mechanism to protect against desiccation by evaporation. In A. rebeli septum formation might be either an autonomous development or triggered by a specific, relatively high permeability of the uppermost whorls. In A. spratti it might be in- dicative of repair, resulting from an accidentally dam- aged apical portion of the shell. Relatively thin apical whorls, due to partial secondary shell thickening below a septum, and often resulting in decollation, also occur in Perricria dausiliaeformis Tap- parone Canefri, 1878 (Figures 6,7) and Taheitia hein- richi Gittenberger, 1989 (Figures 8-10), two terrestrial snail species from New Guinea. They belong to the fam- ilies Megaspiridae [after Zilch, 1959:376] or Coelociidae [after Nordsieck, 1986:111] (Pulmonata) and Truncatel- lidae (Prosobranchia), respectively. In Perrieria dausi- liaeformis, multiple septa are formed during the lifespan Page 40 THE NAUTILUS, Vol. 109, No. 1 of an iiulivickial, resulting in repetitive decollation, as in Ruttiina decoUata. We studied adult, but iicit fully de- collated specimens of both species, in which shell whorls were present above the septum. Extremely thin shell walls were observed just above the septum, suggesting that local resorption had occurred there prior to septum formation (Figures 7,10). In view of the low number of gastropod shells studied with X-rays, we may expect that the decollation mech- anism reported here might be more widely spread among gastropods. St\ lciiiiiiiatii|)ti(ira), Archi\ liir Mdilu.sl^t'nfciinde 106:119- 121. Nordsieck, H 1986. The system of tlie St\ lomniatophora (Gas- tropoda), with special regard to the s\stematic position of the C:laiisiliidae, II Importance of sliell and distriljution. Archiv fiir Mollusling on conspecific eggs; (3) lar- vae and newK hatched snails pre\ ing on conspecific eggs' (Baur, 1992: 103). In that re\iew, Baur described can- nibalism in the terrestrial slug Deroceras laeve (Miiller, 1774) as adults preying on conspecifics, citing Rollo and Wellington (1979). However, although describing it as a very aggressive species, Rollo and Wellington (1979) made no remark on cannibalism in Deroceras laeve. Anecdotal notes similarly seem to suggest that cannibalism in Der- oceras laeve occurs as adults or juveniles cannibalizing other adults and juveniles. For example, Karlin and Nae- gele (1960:14) noted that "it was impossible to hold more than a few specimens of laeve in a rearing cylinder since they exhibited strong cannibalistic tendencies" and that "on several occasions, an indi\idual of this species was actually seen to attack and feed upon a smaller slug of the same species". Pillard (1985:14) also obser\ed "mem- bers of Deroceras laeve feeding on conspecifics". During the past few years, while raising a large num- ber of Deroceras laeve in the laboratory for other pur- poses, I observed numerous cases of cannibalism in this species. In this note, I describe these observations with respect to Baur's three categories noted above. I. Adults or juveniles cannibalizing other adults or juveniles. Although this is the form of cannibalism that has been described by previous authors as occurring in Deroceras laeve, I observed it only once when one larger juvenile (extended body length ^ 17 mm) killed a much smaller juvenile (extended body length ~ 8 mm) and subsecjuently consumed most of the victim. There was no food available in the container. As suggested by Karlin and Naegele (1960), this form of cannibalism may be due to overcrowding or an inadequate diet. On the other hand, I also observed numerous attacks, sometimes re- petitive attacks, among adults and juveniles held in the same container, supporting previous observations by Rol- lo and Wellington (1979) that Deroceras laeve is very aggressive. However, such aggression rarely led to can- nibalism since most victims were able to escape the at- tacks very quickly. While high proportions (as high as 56%) of slugs collected from the field were observed with severe wounds, I do not interpret these as indications of cannibalism in natural populations of this species. 2. Adulls (and juveniles) cannibalizing eggs. In captiv- ity, this typically occurs when the diet is nutritionally unbalanced or when food is not available. Most slugs described here were raised in petri dishes with 3-5 mm of potting soil on the bottom. L nder such rearing con- ditions, adults or juveniles rarely consumed eggs present, provided that food (fresh lettuce) was available. Only on a few ( <5) occasions, was a slug found to have consumed one of the eggs it just laid w hen plenty of fresh food was present. On the other hand, when slugs were raised in petri dishes lined with only wet Kimwipes, 14 of the 16 slugs observed for longer than 3 weeks consumed pro- vided eggs as well as lettuce. Juveniles that ate both eggs and lettuce grew much faster than those that ate only lettuce (unpublished data). 3. Halchlings cannibalizing eggs. This is the most com- mon form of cannibalism that I observed in Deroceras laeve. I disco\ered that newK hatched slugs (1-3 weeks old) of field collected parents often ate their sibling eggs. I also observed this in the hatchlings of laboratory raised, isolated slugs. Egg cannibalism by Deroceras laeve hatch- lings occurred regardless of rearing conditions. The pro- pensity for cannibalism seems to vary both among pop- ulations and among individuals from the same popula- tion. Some hatchlings appeared to prefer eggs to fresh lettuce. Of 25 hatchlings supplied with both eggs and lettuce, four consumed eggs first. Others readily con- sumed both eggs and lettuce. In an experiment where hatchlings were fed only with eggs, however, 5 of a total of 23 hatchlings died after 5-14 days without eating an egg. While it is unclear whether trophic or nurse eggs exist in Deroceras laeve, cannibalistic hatchlings often consumed eggs that did not have a developing embryo. A few hatchlings consumed eggs that had early devel- oping embry OS. Contaminated or infected eggs were nev- er consumed. Cannibalistic hatchlings of Deroceras laeve display a typical behavioral pattern when consuming eggs. They first break the outer coat of the egg, then eat their way into the inside of the egg and gradually eat part or all of the egg from the inside out. Sometimes a hatchling moves its whole body inside the egg and eats from the inside. It typically takes more than one hour for a new hatchling to consume most of an egg that is about the same mass or slightly larger than the hatchling. A can- nibalistic hatchling may abandon an egg at any time. Page 42 THE NAUTILUS, Vol. 109, No. 1 Table 1. Weight comparison between cannibalistic and non- cannibalistic hatchlings of Deroceras laeve. Means and standard deviations (in parentheses) of the weight at hatching and the weight one week after hatching are presented along with the difference between the two measurements (i.e.. weekly growth rate). Non- Cannibal- cannibal- istic istic T-test (n = 1.5) (n = 8) p- value Weight at hatching (mg) 1,240(0.269) 1.388(0.247) 0.2130 Weight after a week (mg) 5.507(2.514) 3.325(1.536) 0.0369 Difference (mg) 4.267(2,344) 1,9.38(1,400) 0,0182 especially if it is disturbed during the beginning or end- ing stages of egg consumption. DISCUSSION Egg cannibalism by hatchlings is the main form of can- nibalism in Deroceras laeve. Adults and juveniles can- nibalizing other adults and juveniles or eggs may occur, but probably only under severe conditions. As described above, egg cannibalism by hatchlings does not seem to be induced by abnormal conditions. Therefore, it could be common in natural populations. In species such as Deroceras laeve, however, it may be difficult to observe egg cannibalism by hatchlings in the field. Eggs of Der- oceras laeve are not only small (mean ~ 3 mg) but also typically buried in the soil or laid beneath boards and other objects, and hatchlings usually do not emerge to the surface immediately after hatching. Egg cannibalism by hatchlings is also known in several other terrestrial gastropods (Baur 1992). For example, newly hatched snails of Arianta arbustoruni (Linne, 1758) and Helix pomatia (Linne, 1758) feed exclusively on eggs (Baur, 1987a, 1988). Baur (1992) suggested that the age specificity of egg cannibalism in gastropods may be associated with an age-specific occurrence of digestive enzymes. Caruubalistic hatchlings often gain consider- able nutritional and energetic benefits. In Arianta ar- bustorum and Helix pomatia. such benefits are appar- ently so significant that cannibalistic hatchlings do not distinguish between sibling and non-sibling eggs (Baur, 1987b, 1990a, 1990b). My observations indicate that can- nibals in Deroceras laeve enjoy a similar nutritional ad- vantage: cannibalistic hatchlings were more than 1.6 times larger than non-cannibalistic ones a week after hatching, and the growth rate of cannibals was on average 2.2 times the growth rate of non-cannibals. As shown in Table 1, both of these values are statistically significant (p<0.05). It also seems that cannibalistic hatchlings of Deroceras laeve do not distinguish between kin and non- kin eggs (unpublished data). Many questions may be asked about cannibalism in Deroceras laeve. How common is cannibalism in natural populations of this species? How variable is the propen- sity of cannibalism both within and among populations? Does this variation of cannibalistic propensity have a genetic basis? What are the costs and benefits of can- nibalism, and what might be its evolutionary conse- quences? Experimental investigations are needed to an- swer these questions. ACKNOWLEDGMENTS I thank Drs. G. F. McCracken and M. G. Harasewych as well as two anonymous referees for helpful comments on the manuscript. I am grateful to Dr. M. L. Pan for providing laboratory space. Financial support was pro- vided by the Theodore Roosevelt Memorial Fund of the American Museum of Natural History and Sigma Xi, the Scientific Research Society. LITERATURE CITED Haur, B. 1987a. Effects of early feeding e.vperience and age on the cannibalistic propensity ot the land snail .\riunta arhu.sluruin. Canadian Journal of Zoolog) 65:.3068-.3070. Baur, B 1987b Can cannil>alistic hatchlings of the land snail Arianta arlmsturum distinguish between sib ami non-sib eggs? Behavior 103:259-265. Baur, B. 1988. Age-specific food preferences in hatchlings of Helix pnmatia (L), Snail F"arming Research 2:14-19, Baur, B 199()a Kgg cannibalism in hatchlings of the land snail Helix puniatia: nutritional adsantage ma> outweigh lack of kin recognition Malacological He\ iew 23:103-105. Baur, B 199()b Possible benefits ot egg canniiialism in the land snail :\rianta arbusturum. Functional Ecologx 4679- 684, Baur, B, 1992, C^annibalisni in gastropods. In: Elgar, M. .\. and B. J. Oespi. (eds.). Cannibalism: Ecology and Evo- lution among Diverse Taxa Oxford I'niversity Press. Ox- ford. England, p. 102-127. Elgar, M. A. and B J Crespi (eds ) 1992. Cannibalism: Ecol- ogy and Evolution among Diverse Taxa Oxford I'niversity Press, Oxford, England. 361 pp Fox, L. R, 1975, (Jannibalism m natural [xipulalions ,\nnual Review of Ecology and Systematics 6:87-106, Karlin, E. J. and J. A. Naegele. 1960. Biology of the Mollusca of greenhouses in New York state. Cornell LIniversity Ag- ricultural Experiment .Station Memoir 372:1-35. Pillard, D A 1985 Xhicus trail following by the slug Der- oceras lacvc (Miiller) Journal of the Teimessee .Academy of Science 60; 13- 15. Polis, G 1981 The evolution and dynamics of intraspecific predatit)n. ,\muial Review ol licology and S\'Steniatics 12: 22.5-251, Rollo, C, D, and W (;, Wellinglon, 1979, Intra- and inter- specific agonistic behavior among terrestrial slugs (Pul- monata: St\ lininalophora) Canadian Journal ol Zoology 57:846-855, INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the hiolog)' and svsteinatics of moUusks. Maiiuseripts de- scribing^ original, iinpnhlished research as well as review articles will be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not re- (jnire an abstract. Notices of meetings and other items ol interest to malacologists will appear in a news and notices section. 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Box 7279 Silver Spring, MD 20907-7279 THEt^NAUTILUS CONTENTS Volume 109, Numbers 2 and 3 October 12, 1995 ISSN 0028-1344 Gary A. Coovert Holly K. Coovert Revision of the Subraspecific ClassiBcation of Marginelliform Gastropods 43 Marine niological Lsiioratory/ WoofJs Hol« Oceannnraphic Instilutian OCT;- ij. MA 02543 THE NAUTILUS 109(2 & 3):43-110, 1995 Page 43 Revision of the Supraspecific Classification of Marginelliform Gastropods Gary A. Coovert Holly K. Coovert Dayton Museum of Natural History 2600 DeWeese Parkway Davton, Ohio 45414 USA ABSTRACT The supraspecific classification of marginelliform gastropods, formerly treated as the single family Marginellidae, is revised This revision is based on all available information for all Recent species plus a representative sampling of the fossil fauna, and includes all Recent and fossil genus-group names. All published information is summarized, and additional dissections, concen- trating on the foregut and radular morphology, are reported, nearly doubling the number of species known anatomically Fundamental differences between two groups of marginelli- form gastropods are discussed. Cystiscids have internal shell whorls partially resorbed and columellar plications reduced internally. Their radular morphology differs fundamentally in having narrower, more numerous rachidian plates, a different subradular membrane structure, separate odontophoral carti- lages, and in the plate development on the posterior end They differ anatomically in tfie possession of a typical neogastropod radular sac. True marginellids have unmodified internal whorls and columellar plications continuous internally. Their radulae are shorter and broader, odontophoral cartilages are fused an- teriorly, and a marginellid buccal pouch is present. The Oli- vidae have the same specializations as the cystiscids and these two groups are considered to have a common ancestry. On this basis, the Cystiscidae is recognized as a valid family and not considered to be closely related to the Marginellidae. The an- cestry of the Marginellidae is discussed and the family is con- sidered to be related to the Volutidae. The Cystiscidae is divided into four subfamilies, three of which are described as new Nine Recent genera, plus one fossil genus, are recognized. Plesio- cystiscus. new genus, is described on the basis of a triserial radula and other primitive characters The Marginellidae is divided into two subfamilies and three tribes, two of which are new. A total of 22 Recent genera are recognized in this family, plus nine fossil genera. Serrataginella. new genus, is described on the basis of its unique radular and shell morphology A systematic section lists synonyms and type species of each genus, along with a diagnosis, description, and summaries of the dis- tribution and fossil record. The phylogeny of both families is discussed, and keys to Recent genera in both groups are pre- sented. Key ivords: Cystiscidae, Marginellidae, marginelliform gastro- pods, systematics, supraspecific classification, new genera, new subfamilies, anatomv INTRODUCTION The best classifications are thus based on the widest pos- sible variety of comparative data. Richard E. Blackwelder (1964;25) The higher classification of the family MarginelUdae has long been in a state of confusion. Especially useful for an overview of the taxonomic history are the papers of Coan (1965) and Coomans (1976). The early mono- graphs of G. B. Sowerby II (1846), Reeve (1864-1865), Weinkauff (1878-1879), and Tryon (1882-1883), as well as Tomlin's (1917) catalog, treated all members of this family under the single genus Marginella. Many popular works still treat all of them under this single genus, while others use generic names inconsistently or erroneously, basing them primarily on superficial similarities of the shell. Coan's (1965) important reclassification was a ma- jor improvement, but is now outdated by new infor- mation. The confusion over the classification stems from the fact that shell characters alone were used in the early classifications, or were largely used in later classifications. Although many good conchological characters exist, they have generally been misinterpreted, or, in some cases, unrecognized. Information on radulae and external anat- omy was widely scattered in the literature, and internal anatomical data, only recently available, is represented by a very small sample. The senior author has spent the last 13 years working toward a revision of the supraspecific classification of the family Marginellidae on a world-wide basis, studying all aspects of these animals, including shell characters, ex- ternal anatomy, radulae, internal anatomy, and obser- vations of live animals, including egg capsules. During this process, many new characters were discovered. We feel enough information has finally been gathered that a stable supraspecific classification can be presented, summarizing this research and presenting a complete classification, including complete synonymies of all Re- cent and fossil genus-group names, along with an inter- pretation of the phylogeny of this group. Page 44 THE NAUTILUS, Vol. 109, Nos. 2 & 3 MATERIALS AND METHODS Since most species were originally described on the basis of the shell, this character complex is of primary im- portance. Because the eventual aim is to completely monograph this group at the species level, initial work involved acquiring every original description for each Recent species named, as well as all significant subse- quent descriptions. A comprehensive species list was compiled from the main catalogs of the family (i.e. Red- field, 1870; Tomlin, 1917; Wagner & Abbott, 1978) and all issues of The Zoological Record and other standard nomenclators were consulted, resulting in the publication of a comprehensive bibliography (Coovert, 1988e). This body of information primarily dealt with the shell, and much effort was expended on studying and analyzing the conchological characters of all Recent species (Coo- vert, 1988d). The type species of all genus-group taxa were closely scrutinized (Coovert, 1986a, 1986b, 1986d, 1986e, 1987a, 1987c, 1987d, 19871, 1987h, and 1988b). A vast amount of information was accumulated, includ- ing the original descriptions of every nominate species, most subsequent descriptions, a large number of pho- tographs of type specimens and other material, drawings, tabular sheets recording locality data and shell measure- ments for thousands of specimens, keys to species groups, graphs of morphometric shell data, comparative charts, and personal observations and notes. All taxa were pro- visionally determined to be valid, synonymous, or un- described, and subsequently assigned to a genus. Further research refined these synonymies and generic alloca- tions. From this large database, a comprehensive species catalog of every valid Recent species was compiled, bas- ing all generic allocations and synonymies on the work outlined herein, not on previous assignments. The generic descriptions, diagnoses, and species counts contained in the systematic section of this paper are summarized from this database. A total of 766 valid Recent species, in- cluding 77 undescribed species, is currently recognized for this entire group. Literature on fossil species was not as comprehensively studied as that on the Recent fauna, but the original descriptions of all genus-group names were studied and a clear determination of each type species was made. A large sampling of additional literature on fossil species was obtained and studied, representing all regions world- wide. Specimens and descriptions of fossil species from the western Atlantic were particulary well represented. Generic allocations of fo.ssil species are based on our own work, and do not necessarily agree with previous assign- ments. Our approach to the fossil fauna is conservative. Only specimens or literature personally examined were included in our database. A good representation of the fossil fauna has been obtained, and is presumed complete as far as genus-group names are concerned. During this 13-year period, shell specimens from a wide variety of .sources, including a large number of museums and individuals (.see acknowledgements), were studied, largely concentrating on the Recent fauna. Shell measurements, including maximum length par- allel to the axis and maximum width perpendicular to the axis, were made with a Spencer AO binocular dis- secting microscope using a calibrated optical reticle for minute to small shells under 5 mm. For larger shells, an electronic digital Max-Cal caliper was used with a res- olution to 0.01 mm. Internal plications were initially studied by noting internal features on shells with gastro- pod drill holes or breakage. More detailed studies were conducted by grinding a hole on the ventral surface of the shell with a fine metal file. Some shells were trans- versely sectioned with a fine jeweler's saw, ground to a smooth surface on a fine-grit whetstone, then polished on a glass plate with toothpaste as a fine abrasive. Methods used for the extraction, illustration, and anal- ysis of radulae are given in complete detail in Coovert and Coovert (1987) and only summarized here. Most radulae were extracted without damaging the shell by soaking and flushing the shell with a solution of 10% potassium hydroxide (KOH) using a micropipette. After rinsing with acidic water, radulae were transferred to a microscope slide and mounted in glycerin jelly. Drawings were made using an optical reticle grid in a standard light microscope. Radular data, reported for each genus in the systematic section, includes a shell length to rad- ular width ratio plus the Radular Index. The Radular Index is derived by dividing the number of rachidian plates by the number of cusps per plate, and requires a complete radular ribbon, w hich was not always available. Routine extraction using KOH destroys many details of the delicate subradular membrane, especially in true marginellids. For a detailed study of these features, rad- ulae were removed during dissection without the use of KOH and studied in glycerin before mounting (see be- low). Study of the developing posterior end of the radula was made from previously mounted material. All per- sonally prepared material, totalling 224 specimens of 97 species, was mounted using glycerin jelly and studied with a standard microscope. Through the courtesy of Dean Hewish, SEMs of radulae from 99 specimens of 67 mostly Australian species, with complete locality, shell, and morphometric data, were made available. A review and summary of published information is given in Coo- vert (1989b), with additional radular data presented in Coovert and Coovert (1990). A combined total of 190 radulate species, plus an additional 30 species known to lack a radula, have been studied in detail. The external anatomy from published sources was summarized in Cloovert (1987g). Many photographs and drawings, provided by colleagues, were also studied. Live material was examined using a Spencer AO binocular dissecting microscope, then carefully drawn in color. In more remote localities, a Rupen wide-field 16 X hand lens was used. Photographs were taken w hen possible. Data on the external anatomy of 183 species is sum- marized in this paper. All published data on the internal anatonn was studied in great detail. (Charts of potentially useful taxonomic characters of all 13 anatomically known species were G. A. Coovert and H. K. Coovert, 1995 Page 45 constructed, patterned after those in Harasewych and Kantor (1991). Using this information as a guide, and carefully choosing taxa to augment this published data, dissections were completed on 11 additional species, plus one previously reported species that lacked sufficient data. We concentrated on features of the foregut, as these were determined to be of most value taxonomically . Specimens were dissected using a Spencer AO binocular dissecting microscope. Gross dissections were completed on mul- tiple specimens of a species when available, with a total of 26 specimens being dissected (Table 1). Detailed, an- notated drawings were made during all stages of dissec- tion. Preservation varied among material due to a wide range of sources and preservatives. Larger animals were extracted from shells with a fine jet of water after briefly soaking them in liquid dish soap. Smaller shells were first measured, then cracked using a small hand vice. Odon- tophoral cartilages were carefully extracted from the buccal mass of radulate species, manually cleaned of adherent tissue, and mounted on microscope slides using glycerin jelly as a mounting medium. The subradular membranes of these radulae were cleaned of extraneous tissue after extraction and studied in glycerin prior to permanent mounting in glycerin jelly. Potassium hy- droxide was not used in the preparation of these carti- lages or radulae at any stage. A list of the dissected material, with locality data, shell measurements, and oth- er details, is presented in Table 1. The following revision is based on all available infor- mation of every Recent species plus a representative survey of the fossil fauna. Most importantly, the type species of all marginelliform genus-group names, both Recent and fossil, were studied and fully clarified. Gen- era are based on groups of related species, not just their type species. A very conservative attitude was taken to- wards recognition of higher taxa. Shared, derived char- acters were searched for to unite groups rather than using minor differences to split groups. The rank of subgenus is not utilized in the classification presented as this would require additional research and necessitate formal rec- ognition of additional taxa. We prefer the useage of informal "species groups." The proposal of new taxa has been avoided unless strongly supported. ABBREVIATIONS AND CONVENTIONS The following abbreviations and conventions are used in the svstematic section and text: =? ? t t aff. AMNH AMS synonymy questioned (species) synonymy questioned (genus) (preceding genus), denotes extinct genus (following species), denotes fossil species affinis. having affinity with but not identical with American Museum of Natural History Australian Museum, Sydney ANSP Academy of Natural Sciences, Philadelphia BM(NH) British Museum of Natural History DMNH Delaware Museum of Natural History GAG collection of Gary A Coovert ICZN International Code of Zoological Nomencla- ture M original designation by monotypy MHNG Museum d'Histoire Naturelle, Geneve MNHP Museum national d'Histoire naturelle, Paris NMV National Museum of Victoria OD original designation OD (M) original designation as type species, but also monotypic OD (T) original designation as type species, but also tautonomous SD subsequent designation as type SD (M) subsequent designation as t\pe species, but also monotypic sic to signify exact transcription T original designation by tautonomy TS type species USNM National Museum of Natural History, Smith- sonian Institution In the systematic section, the following sequence is used for each generic synonym: Genus in original combination, followed by author, year, page number(s) TS as cited by original author of genus (or if not originally cited, as cited by author of SD) TS in original combination of author of species, if dif- ferent from above Senior synonym if applicable Designation of type species (i.e. M, OD, OD (M), OD (T), SD, SD (M), T); SD followed by author, year, page number(s) In the systematic section, nomina nulla (misspellings) are not included. Most nomina nuda, unless they have a direct bearing, are also not included. Citing these is undesirable because it imparts a false sense of validity. The comments in Tomlin (1917:242) are applicable. Throughout the remainder of this paper, the term marginellid will be used in the strict sense for the family Marginellidae as defined and restricted in the systematic section herein. The term cystiscid will be used for the family Cystiscidae as defined in the systematic section herein. The term marginelliform will be used in the broad sense of marginellid, equivalent to the traditional concept of the family Marginellidae as conceived by previous authors, and including both marginellids in the strict sense and cystiscids. Terminology pertaining to shell morphology, radular morphology, and anatomy has been used differently and without standardization by previous authors. Consistency and uniformity are important enough to warrant careful selection and concise definition of terms. In addition, several new terms are introduced and defined. Page 46 THE NAUTILUS, Vol. 109, Nos. 2 & 3 Tablo 1. Sex, voiu-licr niiiiiher, localit) data, coniment.s. and shell measurements of disseeted material All dissected specimens, incliidini; shells and slide preparations, lia^c been retained in the collectidii of (;ar\' A. (-oovert. Shell Species Sex Vouchers Localit v/C'omments Length Width (mm) (mm I Bullata Imllata (Born, 177S) male Dt'titintargo elninwola iC<.>u- rad, 1834) female Dentinmrgn cbiirncola (Con- male rad, 1834) Hyalina pallida (Linne, 1758) male Marginella glabella (Linne, lemaie 1758) Marginella sebaitiani Marche- male Marchad & Rosso, 1979 Marginella sebastiani female Persicula interruplolineata male (Megerle von Miihlfeld, I81(i) Persicttia intcrrupiulineala Persicula interrupt vlineata Persicula interruptolineata Persicula interruptolineata Persicula masirana Roth ii Pet- it, 1972 Prinmnt aff aletes Rolh, 1978 male .Note 2 Prunum aff. aletes Prunum aff. aletes Prunum guttalum (DilKwn 1817) Prunum gutlatum Prunum gutlatum Prunum prunum ((JmeliM, 1791) letnalc GAC Acc. #I(> 90 GAC 1177A GAC 1448 GAC Acc. #4- 94 C GAC M2548 GAC: M2549A GAt; M2549t; GAC 1581 A lemaie GAC 1581B female GAC 1581C female GAC 1581 D female GAC 1581 E 2 males GAC Acc. #4 94 A, B male GAC 1518A female GAC 1518B male GAC 1518C male GAC 89 Acc. #2 male GAC 1172A (;AC 1 17(IB (;ac: 1.582 41.8 38.8 39.8 14.7 23.8 21.4 23.4 9,6 Baliia de Todos Santos, Bahia State, Brazil, 5 52.1 27.8 meters depth; ex. E. C". Rios; ex. isopropyl alcohol; Note 1 Stock Island, on N. side of U.S. Rt. 1, E. end 6.9 3.1 of island, Monroe Co., Florida, under algae covered rocks in 0.3 m, Sept. 7, 1984, col- lectors G, A & H, K. Coovert; ex. isopro- pvl alcohol Stock Island, on N, side of L'S Rt, 1, E, end — — of island, Monroe Co,, Elorida, hand- dredged in sea grass beds in 0,3 m, Nov, 30, 1988, collectors G, A, & H, K, Coovert; ex, isopropyl alcohol (no data); ex. Robert Lipe; ex. ethyl alcohol, — — animal only "Banco Canario-Saharia,' (apparently a fish- ing bank between Spanish Sahara and the C^anary Islands), March 20. 1991; ex Jose Aguiar Morales; ex. methyl alcohol, subse- quently softened with trisodium |)hosphate (same data as above) (same data as above) N. end of Playa La Galera, Isia de Margarita, Venezuela, NE side of ba\ , hand-dredged on sandy mud with small patches of Turtle Grass, Oct, 22, 1992, collectors G A, & H K, Coovert; ex, isopropyl alcohol (same data as above) (same data as above) (same data as above) (same data as above) Masirah Island. Oman, shallow water, low — — tide, collector Donald T Bosch, ex Robert Lipe; ex. eth> 1 alcohol; animals only small island just off Tamarindo Beach, S. of 18.8 12.1 Tamarintlo Diria Hotel, CJuanacaste Prov- ince, Costa Rica, under rocks, low tide, Oct. 18, 1989; collector Peter Bright; ex. isopropyl alcohol (same data as above) 19.4 12.2 (same data as above) 20.4 13.4 •Airport Reef, I'tila Island, Bay Islands. Hon- — — duras, shallow water under rocks; ex Rob- ert Lipe, ex, isoprop\ 1 alcohol, animal oiiK chamiel behind Lagoon Motel, Marathon, 16,5 9,7 Key Vaca, Monroe Co , Florida, under rocks, snorkeling in 0-1 m, Sept. 5, 1984, collectors G. A. it H K Coovert; ex. iso- propyl alcohol (same data as above); Sept 1, 19S1, late juve- 14.8 8,0 iiile Mal;i Redonda, Isla de Margarita, Venezuela, 30,7 15.8 shallow sand and mud flats with scattered Turtle (irass. low tide, (Kt 24, 1992, ex John Wolfe; ex, isoprop\ 1 alcohnl 12.5 8.4 12.2 7.9 11.6 7.5 13.5 8.4 G. A. Coovert and H, K. Coovert, 1995 Page 47 Table I . ( 'oiiliiiiifil Sex Species Pri/num rooscvciti (Bartscli ^ male Rehder, 1939) Note 3 Pruninu rooscvciti male Volvarinu aft (ucnii (Kiener, male 1834) Note 4 Voloariiui aft dvcna lemale Volvarinu aft menu lemale Shell N'ouchers .ocality /Comments Lengtli Wkltli (mm) (mm) #26- Walkers Clay, Batiamas, snorkeiing at 1 m, — — grass and algae, night, June 17, 1988, col- lector Robert Lipe; ex. isopropyl alcohol; animal only #2(> (same data as above) 16,9 1 I 0 15 channel behind Lagoon Motel, Marathon, 11.3 5.0 Key Vaca, Monroe Co., Florida, under rocks, snorkeiing in 0-1 m, Sept. .5, 1984, collectors G. A & H. K. Coovert; ex iso- propyl alcohol G.^C 1170C (same data as above); Sept. 4. 1984 11,3 .5,0 G.'\(: 1170E (same data as above); Sept, 4, 1984 10 1 4 5 GAC: Ace 88A GAC; Ace 88B GAC 1172 Note 1: Radula figured in Coovert and Coovert (1990:2, fig. 1). Note 2: Shells were significantly larger and much more brilliantly and differently colored than the nominate deep-water form (from 3 to 82 m). More research is necessary to establish the status of this taxon. Note 3: The radula of this species is closest to P. carneum (Storer, 1837), the animals greatly differing in the external coloration. The radulae are very distinct from the conchologically very similar Prunum amabile (Redfield, 1852), also from the Bahamas, whose radula was described in Coovert and Coovert (1990:5-6). Note 4: See comments in Coovert and Coovert (1990:35) regarding the status of the Florida form of this species. COMPARATIVE ANALYSIS OF SHELL MORPHOLOGY Contrary to the opinion of many previous authors, there are many conchological characters that are very useful in the classiBcation of margineliiform gastropods. Those most useful for higher categories are emphasized. External varix: The external varix or "margin," for which these shells are named, is one of the more im- portant shell characters. Other authors have used the terms "margin," "marginal varix," "varix, "labial varix," "outer varix, ' or "varical callus, " not clearly differen- tiating between a true external varix and a merely in- ternally thickened lip. The term external varix is herein used in preference over the above terms as it more clearly makes this differentiation. A shell with an external varix has a weakly to distinctly raised margin of callus depos- ited externally along the labial edge, easily seen by close dorsal examination of the shell. In all marginelliforms, the outer lip is at least some- what strengthened or thickened internally in the adult, but many genera completely lack an external varix. This fact has received very little attention in the literature and most original descriptions fail to mention the pres- ence or absence of this important character. Shells lack- ing a varix are even and smooth along the external labial edge, without a discernable raised margin of callus. The outer lip in such shells is usually marked by a change in angle when viewed from the apex and is often a different color. Presumably, a strengthened lip, either internally, externally, or both, imparts adaptive value in protecting the snail from predation. Most cystiscid genera, including "Cystiscus" (the group with a triserial radula), Cystiscus s. str., Crithe. Gib- berula, and Canalispira, completely lack an external varix. Only the cystiscid genus Persicula varies in this character, with some species possessing a well-developed e.xternal varix, whereas others completely lack it. Gran- ulina and Piignus are the only cystiscid genera that al- ways possess a strong external varix. Because the very strong axial costae in Extra resembles varices, the pres- ence of this character is indeterminate. Based on other conchological characters, Extra has been placed with Crithe and Cystiscus, both non-varicose genera. Marginellid genera, as opposed to cystiscid genera, nearly always possess a strong external varix. Hyalina, a notable exception, usually lacks an external varix. The genera Prunum and Volvarina vary in this character, some species possessing a strong varix, some a weak varix, and other species completely lacking it. The remaining marginellid genera always possess at least a weak external varix. This usually very strong, distinct varix often con- tinues around the anterior edge of the siphonal canal. A few species have an extremely thickened external varix that is doubled or "duplicate." Ontogenetic lip development, from initial formative stages in the juvenile, through subadult, to the fully adult stage, has seldom been reported in the literature. We have studied late juvenile and subadult shells for many years in an attempt to ascertain stages of labial devel- opment. The external varix is initially formed by an outward upturning of the thin lip edge in most species with a varicose adult shell. Callus deposition subsequent- ly occurs internally and externally, resulting in the thick- Page 48 THE NAUTILUS. Vol. 109, Nos. 2 & 3 ened, adult varix. We have observed this sequence in the cystiscids Persicula cingulata (Dillwyn. 1817) and P. persicula (Linne. 1758), and in the marginellids Erato- idea margarita (Kiener, 1834), Marginella rosea La- marck, 1822, and Glabella adansoni (Kiener, 1834). Be- cause the situation in Prunum and Volvarina is more complex, we transversely sectioned, polished, and studied many shells from these genera. Preliminary results in- dicate that some varicose species of Prunum, but not Volvarina, form the varix as described above. This can often be seen in cross-sections, with the initial varix and subsequent concentric growth rings showing clearly. Other varicose species of both Prunum and Volvarina initially curve the thin, fragile, developing lip edge in- wardly, subsequently adding further callus deposits in- ternally to strengthen and thicken the edge. Only later are significant external callus deposits added in those species with a varicose adult shell. Preliminary results with shell cross-sections of Prunum and Volvarina in- dicate a difference in how this occurs. Varicose Prunum species with an initially incurved lip produce the varix by adding shell material to both surfaces while extending the growing edge. This can be seen in cross-section as very fine concentric growth rings centered around an initial growing point. Varicose Volvarina species, on the other hand, produce the varix by callus deposition in- ternally and externally, without much elongation of the labial edge and lack obvious concentric growth rings. The same difference appears to exist between non-var- icose Prunum and Volvarina species, but the external varix simply does not form in these taxa. Additional study of this character is needed before definite conclusions can be drawn, but preliminary results indicate the ex- istence of a potentially useful taxonomic character. Labial Denticles: Many marginelliform shells have an outer lip with distinct denticles, referred to herein as labial denticles or denticulate lip. This is preferred over such terms as "teeth," "crenulations,"or "crenate." Pres- ence or absence of labial denticles is sometimes of generic value but is usually only of specific value. Lirae are a different type of labial structure and are very useful taxonomically. They are usually erroneously referred to as "denticles" or "teeth" in the literature, but the dif- ference is very important. Denticles are small bumps or tooth-like projections on the inside edge of the outer lip and generally do not extend into the aperture. Lirae are thin spiral ridges on the inside of the outer lip oriented perpendicularly to the lip edge and extending well inside the aperture. They resemble denticles only at the outer edge of the lip. The only marginelliforms possessing lirae (but not in all species) are the cystiscid genera Persicula, Canalispira, and Gihbertda. These lirate genera usually lack an external varix. Columellar Plications: Another extremely important character complex is the number and formation of col- umellar plications. The terms "fold," "plica," and "plait," used by other authors, are considered inappropriate for a spirally wound ridge of shell material that is di'posited upon the columella and is not an actual fold of shell material. The term columellar plication is thus preferred. The anteriormost plication of marginelliform shells does not differ fundamentally from the adjacent plication, and is thus not differentiated by a separate term, such as "siphonal fold or "basal told." This anteriormost pli- cation is herein considered the first plication, thus count- ing anterior to posterior. In many animal groups, serial features are likewise counted in a standard anterior to posterior direction. A few authors, mostly of older works, numbered plications from posterior to anterior, resulting in the anteriormost plication being variously numbered. A survey of marginelliform gastropods reveals that the first columellar plication borders the anterior edge of the columella in all cases (usually continuing and merging with the external varix) except Afrivoluta, which has a very volute-like columella. Two fundamentally different plicational conditions occur in marginelliform species. These are correlated with the type of internal whorls present. Internal Whorls: One group has typical neogastropod internal whorls that are simply previous body whorls subsequently enclosed by additional whorls without fur- ther modification. The columellar plications remain un- modified for their entire length. This type of plicate internal whorl has also been observed in the families Turbinellidae, Cancellariidae, Mitridae, Costellariidae, and Volutidae. The internal whorls in true marginellids are maintained at their original thickness with no ap- parent resorption or further modification and are herein termed unmodified internal whorls (figure 1). Taxa with unmodified internal whorls have columellar plications remaining at their original strength and number through- out subsequent whorls. These plications can be found intact internally upon the entire length of the columella. They are here termed continuous columellar plications and are normally oriented nearly perpendicular to the shell axis. The usual number of plications in this group is four, but varies from two to six among different species, although not normally varying within a species. In those species with five or six plications, the fifth and sixth continue internally for at least a hall whorl, and often are fully continuous. These are also considered contin- uous plications in this paper. In some species, a weak "false fifth plication" (or less commonly a "false fourth" or "false sixth ) is present. This may be either a parietal lira that does not continue into the aperture for more than an eighth of a whorl, or a denticle, but neither are true plications because they are not continuous. Within marginelliform gastropods, unmodified internal whorls and continuous columellar plications are found only in species herein assigned to the Marginellidae s. str. Another group has extremely thin, largely resorbed internal whorls, olten partially missing due to complete local resorption or breakage. Proceeding inward, the first three (anterior) columellar plications in the aperture are quickly reduced to two, the third simply diminishes with- in a half whorl internally. The remaining two are further reduced to a single, sharp, very oblique columellar edge G. A. Coovert and H. K. Coovert, 1995 Page 49 Figures 1-4. Internal shell vvhorls. 1. Prunum priinum (Gmelin, 1791), showing unmodified internal whorls. GAC, no data Length 30.9 mm 2. Persicula persicula (Linne, 1758), showing cystiscid internal whorls. GAC Ace. # 54-87, Joal River, Senegal, on sand flats at mouth of river. Length 20.0 mm. 3. Plesiocystiscus jewettii (Carpenter, 1857), showing cystiscid internal whorls. GAC Ace. # 14-90, California. Length 5.7 mm. 4. Cranulina hadria (Dall, 1889), showing modified cystiscid internal whorls. GAC 824, Tampa Bay, Pinellas Co., Fla.. 0.3 — 0.6 m, shallow grass flats. Length 2.0 mm. within one full internal revolution. This single, sharp columellar edge essentially becomes the axis of the shell. The shape of the internal whorls are very different from the original juvenile body whorls that were subsequently resorbed after being enclosed by the next whorl. Original juvenile shell color, patterns, and apertural columellar plications, generally more than two, are all completely resorbed. Among marginelliform gastropods, this type of internal whorl is found only in species here assigned to the Cystiscidae. They are here termed cystiscid internal whorls (figures 2, 3). Columellar plications in species possessing cystiscid internal whorls differ fundamentally from the continuous plications found in species with un- modified internal whorls, primarily because they are quickly reduced to a single columellar edge, and are not continuous internally. They are here termed internally reduced columellar plications. Posterior to the three in- ternally reduced columellar plications are one to ten parietal lirae superficially resembling columellar plica- tions. These were not differentiated from columellar pli- cations by most previous authors. Parietal lirae are some- what smaller and weaker than the three anterior plica- tions, do not continue into the aperture for more than a quarter turn, and gradually diminish posteriorly. This results in a total of up to 13 "plications," of which only the anterior three are considered true, but highly mod- ified, plications. This columellar arrangement, composed of internally reduced columellar plications plus parietal lirae, is herein termed multiplicate. This term has been used by some previous authors, but was not clearly de- fined. In a few cases, there may be parietal denticles posterior to the parietal lirae, but these are even shorter. A very few species of Cystiscus have only two columellar plications (one species, possibly not a cystiscid, is reported to have only a single plication). A modification of the cystiscid internal whorl has been studied in species herein assigned to the genus Granulina. This modified type is characterized by the first two col- umellar plications (proceeding inwardly) abruptly end- ing within one full revolution internally. At this point, a thin, axially oriented edge begins much as in species with cystiscid internal whorls. However, the two posterior col- umellar plications continue and are situated at the an- terior edge of the more bulbous portion of the mostly resorbed internal whorl. These usually translucent inter- nal whorls are extremely thin and fragile and break very easily. The anterior end of these whorls appears to be open. We believe that this is a modification of the cys- tiscid internal whorl and is herein referred to as a mod- ified cystiscid internal whorl (figure 4). The four colu- mellar plications of Granulina superficially resemble continuous plications, but because the first two quickly terminate, they are considered to be internally reduced columellar plications typical of all other cystiscids. The posterior two continue, and are herein referred to as pseudo-continuous plications because they apparently do not continue for more than a whorl or two internally. Members of the genus Granulina lack the gradually weakening parietal lirae posterior to the columellar pli- cations, and thus they do not have a multiplicate colu- mella typical of other cystiscids. Apertural Characters: The extent to which the aperture is occupied by the columellar plications is useful taxo- nomically. Genera with a multiplicate columella usually have more than half the apertural length occupied by the plications plus parietal lirae. Marginella, Glabella. Dentimargo, and Eratoidea have four strong, continuous columellar plications occupying more than half the ap- ertural length, as do Austroginella and many of its rel- atives. Most remaining typical marginellids have four plications occupying less than half the apertural length. Page 50 THE NAUTILUS, Vol. 109, Nos. 2 & 3 In a few genera, though, the pHcations are crowded an- teriorly. Columellar plications are also very useful ta.x- onomically at the species level. In addition to the number of plications, other useful characters include their height, thickness, shape in cross-section, extent of protrusion from the aperture, their angle, whether they are fused or sep- arate, and whether or not they are excavated inside the aperture. Very little use has been made of these features in the literature. Another feature of the aperture is its relative width, which can often be compared to the lip thickness as a readily available reference. Most genera have an aper- ture that is widest anteriorly, but in some the aperture is broadest medially. Siphonal Notch: The presence or absence of a siphonal notch is an important shell character. Called "anterior notch," "notched anterior canal," or "siphonal canal" by various authors, the term siphonal notch is preferred, as it relates to the function of this feature. A distinct si- phonal notch is present in Persicula and Gibberula, but absent in the related Canalispira. This notch is a typical character of Glabella and most Marginella s. str. The genera Afrivolnta and Marginellona possess a siphonal notch, although it is weak in Marginellona. Austrogi- nella has a distinct siphonal notch, but in the related Mesoginella the notch is very weak to absent (Coovert, 1988b). In all remaining marginelliform genera the notch is very weak or absent. Posterior Notch: A weak to distinct posterior notch is sometimes developed at the junction or commissure be- tween the body whorl and the posterior end of the lip. The posterior end of the lip generally forms an abrupt angle just prior to joining the body whorl, leaving a broad to narrow notch between the lip and body whorl. Less commonly, this notch forms as a weak to distinct groove in the posterior end of the lip itself. Variously termed "posterior notch," "posterior sinus," "posterior canal," "anal canal," or "anal sulcus," posterior notch is pre- ferred, because it is consistent with "siphonal notch." The terms "anal canal" or "anal sulcus" imply a func- tional aspect not borne out anatomically. The presence or absence of this character is often obscure, reducing its taxonomic importance. However, many Persicula, Gibberula, and Canalispira have a distinct posterior notch formed at the junction of the body whorl and posterior end of the lip. The jiosterior notch in Persicula is often accentuated by an adjacent callus deposit on the body whorl. The deeply channeled, narrow notch in Canal- ispira is diagnostic. Protoginella has a posterior notch formed in the posterior corner of the lip itself. Spire: Spire height is often important taxonomically. Several groups have a completely immersed spire and thus an easily recognizable shell sha[)e. In this paper, a low spire is defined as having a length less than 'A the shell length (figures 43, 44, 48, 55). A spire of medium height ranges from V* up to '/i the shell length (figures 57, 73), whereas a tall spire is more than '/i the shell length (figures 49, 70). Spire height often varies consid- erably within a genus and is thus more important at the species level. Spire height is determined by the insertion point of the posterior end of the lip, and can be refer- enced by whether or not the posterior end of the lip joins the body whorl at, above, or below the previous suture. This is a very useful character within Volvarina, for example. Shape: Shell shape is another useful feature, and al- though somewhat subjective, should be carefully de- scribed using standard shape names. Marginelliform shells vary from a nearly round, globose outline to elliptic, obovate, obconic, conic, biconic, subtriangular, pyriform, elongate-ovate, subcylindrical, cylindrical, and other combinations. The shape of the shoulders is very impor- tant in determining overall shell shape, and can be gently curved to strongly rounded or carinate. Size: Shell size is often very valuable as a species-level character and, in a general way, for higher categories. Shell size is here reported as minute for shells up to 2.4 mm in length, small for shells over 2.4 mm up to 6.0 mm, medium for shells over 6.0 mm up to 13.0 mm, moderately large for shells over 13.0 mm up to 25.0 mm, large for shells over 25.0 and up to 50.0 mm, and very large for shells over 50.0 mm in length. Surface Features: Shell texture is another very useful conchological feature. Although most marginelliform shells are perfectly smooth and glossy, some have very characteristic weak to strong a.xial costae. This term is preferred over "axial ribs" or "longitudinal ribs. This character is often useful at the genus or the species level. Pronounced surface texture is very rare in the family, but Pugnus and a few Granulina have a minutely sculp- tured surface. Rivon}arginella species have minute pus- tules ventrally that are part of the callus deposit. Many other species have callus deposits ranging from a thin parietal wash to distinct deposits on the spire, posterior parietal area, or anteroventrally. Ajrivoluta has a very distinctive callus pad (Coovert, 1987a). The placement of callus deposits is species specific and thus diagnostic in the Persicula cornea group (Coovert, 1987b). Many Granulina. Crithe. and other species have a character- istic callus deposit here termed a collabral parietal callus ridge (figures 55, 56). This is a narrow ridge of callus deposited parallel to the lip and usually lying just outside the aperture. It often interesects the columellar plica- tions, causing them to appear excavated. Heavier callus deposits at vulnerable points are interpreted as a pro- tective adaptation against predatory boring. Preliminary observations suggest a strong correlation betw een typical bore hole locations in uncallused species and locations of callus deposits in related species. (]olor: Shell color and pattern are also useful taxonomic characters. Most small specie's are translucent to opaque white. Mo.st larger species are white, but some range from gray or greenish-gray, to brown, amber, flesh-col- ored, yellow, orange, pink, rose, or even red. The base color may he overlain with various pattern elements of G. A. Coovert and H. K. Coovert, 1995 Page 51 a different color. These can consist of spots, irregular blotcfies, spiral lines or bands, axial or oblique bands or streaks, oblique lines, or various combinations of these features. COMPARATIVE ANATOMY OF MARGINELLIFORM GASTROPODS External Anatomy: Features of the external anatomy of 82 species of marginelliform gastropods have been reviewed and summarized (Coovert, 1987g). Since then, a number of additional species have been reported in the literature (Coovert, 1988e:37). Our additional re- search, plus notes and excellent photographs of the living animals shared by others, provided a clearer picture of the external anatomy of this group. Data on 183 species are summarized in this paper. The four basic morpho- logic types are summarized first, along with a discussion of general features of the externa! anatomy. Further details of the external anatomy of each genus can be found in the systematic section. The four distinct animal "types" delineated and sum- marized in Coovert (1987g) are based primarily on fea- tures of the head. The most common marginelliform animal is termed a "Type 2 animal," which has a simple, unmodified head described by some authors as "bifur- cate" (figures 7, 8). The long, slender tentacles emerge from the anterior end of the head, resulting in a bifurcate appearance. Viewed from above, as most photographs are oriented, this bifurcation is not readily apparent. The long, slender tentacles have an eye located on a slight swelling at their bases. The siphon, as in all marginelli- form gastropods, is a simple tube formed by the inrolled edges of an elongated expansion of the mantle edge. The siphon in Type 2 animals is moderately long to very long, and simple or weakly notched distally. A few cystiscid and most marginellid genera have a Type 2 animal, which is considered to be the generalized type. Type 1 animals, represented by Afrivoluta and Mar- ginellona, are characterized by a broad head with a longitudinal medial furrow or channel (figures 5, 6). This channel was described by Harasewych and Kantor (1991: 10) as a "deep tubular channel along dorsal mid-line of head," apparently formed by the ventral fusion of the tentacle bases. These same authors (op. cit., p. 13) noted that this medial channel had not been reported in A/- rivoluta, but our careful examination of a color trans- parency of Afrivoluta pringlei (ex. W. R. Liltved) has revealed that both genera share this feature. Type 1 animals are further characterized by relatively short ten- tacles, a simple, moderately long siphon, and a distinct, subtriangular lateral lappet on each side of the head. A small red eye is present on each lateral lappet in Afri- voluta. Eyes are completely lacking in Marginellona. Type 3 animals are characterized by an elongate head that is longitudinally split dorsally, with the anterior end bifurcate (figures 9, 10). This configuration could be de- scribed as either having the tentacles fused ventrally, resulting in a longitudinal dorsomedial channel, or al- ternatively, as having a longitudinally split head com- pletely lacking tentacles. A detailed anatomical study is needed to clarify this situation. The siphon is either very short and not readily apparent, or completely absent. Eyes are located on the sides of the head, usually in a conspicuous bulge. Type 3 animals are represented by the genus Cystiscus and probably also Crithe. Type 4 animals have a somewhat flattened, longitu- dinally split head, normally with tentacles (figures 11, 12). The anterior half of the head is completely split, usually with the anterior tips pointed and divergent, but capable of closing together. These two lobes are very flexible and capable of much movement. The posterior half is only split dorsally, and fused ventrally. Tentacles, which are rarely absent, are short to long. The eyes are located on the sides of the head, slightly below the ten- tacle bases. The siphon is short to long, simple, and pro- trudes through the siphonal notch at the anterior end of the shell when the animal is active. Type 4 animals are represented by the genera Persicula and Gibberula. Can- alispira animals differ from the typical Type 4 animal in lacking tentacles and a visible siphon (cf. Systematic section for more detail). The siphon of most marginelliform gastropods emerg- es from the left side of the head, but has been described as fused to the head in Marginellona by Harasewych and Kantor (1991;table 3), although these authors stated that the siphon is "fused to head just left of posterior margin of mid-line channel" (op. cit., p. 10). It was described as fused to the mid-dorsal line of the head in Priintim marginatum by Graham (1966:135). All species studied by us (Table 3) have a siphon fused to the left side of the head. There seems to be some difference in interpretation, but probably all marginelliforms have a siphon fused to the head at least slightly left of center. The mantle is another external anatomical feature that is useful taxonomically. Within higher ta.xa, the degree of mantle extension over the external shell surface varies from none to complete coverage. Extended mantles can be smooth, have low, rounded tubercles or pustules, or possess elongate papillae that are either simple or branched. The terms pustulose and papillose are used here. Most marginelliform gastropods have a broad, flat foot, although it may be very narrow in a few species. The relative size and length of the foot may have some value taxonomically. Some species have a medially indented foot, and some (e.g. Hyalina hyalina, H. pallida, and Prunum martini) have an anterior, transverse pedal groove. As far as is known, females have a medial, an- teroventral pedal gland on the sole of the foot. In some Gibberula species, the anterolateral edges of the foot are raised and laterally rolled, forming what Gofas and Fer- nandes (1988:22) described as "parapodia." All these characters may have taxonomic value and need further study. Animal coloration is very useful at the species level, but has limited value for higher classification. Gofas and Fernandes (1988) and Gofas (1989a, 1990, 1992) provid- Page 52 THE NAUTILUS, Vol. 109, Nos. 2 & Figures 5-8. Marginellid external anatomy. 5, 6. Afrivoluta pringlei Tomlin, 1947, e.xample of Type 1 animal. After Liltved (1985) plus pers. obs. of photos (ex. W. R. Liltved), Off Danger Point, western Cape Province, at 2-46 m. Ca. X 0.65. 5. Animal viewed from right side. 6. Dorsal view of head. 7, 8. Prunum aff. aletes Roth, 1978, example of Type 2 animal. GAC 1518, small island just off Tamarindo Beach, S. of Tamarindo Diria Hotel, Guanacaste Province, Costa Rica, under rocks, low tide Shell length 19.4 mm. 7. .'\nimal viewed from right side. 8. Dorsal view of head. Figures 9-12. Cystiscid external anatomy, 9, 10. Cystiscus minutissimus (Tenison-VVoods, 1876), example of Type 3 animal. After Murray (1970) and Coleman (1975). 9. .\ninial viewed from above. Ca. 9.0 X. 10. Dorsal view of head. Ca.l8 X. 11, 12. Persicula interruptohneata (Megerle von Miihlfeld, 1816). example of Type 4 animal. GAC 1581, N. end of Playa La Galera, Isla de Margarita, Venezuela, NE side of bay, hand-dredged on sandy mud with small patches of Turtle Grass. Length 14.7 mm. 1 1. Dorsal view of head. 12. .\nimal viewed from right side. ed color plates of live animals, making excellent use of coloration or "polychromatism" for species differentia- tion. Radula: The radulae of very few species (20 fide Coo- vert, 1989b) were known at the time of the last generic revision of this group (Coan, 1965), Much additional radular information has been subsequently published (see Coovert, 1989b for sutnmary; Coovert and Coovert, 1990). This published data, combined with our unpublished data and observations of SEMs (ex. Dean Hewish), brings the number of species studied to 190. An additional 30 species are known to be non-radulate. Of the 31 Recent genera recognized in this revision, the radulae of only two are unknown. Three basic radular patterns of marginelliform gas- tropods have long been recognized by the senior author. In addition to non-radulate species, the radulate species fall into two different groups. One group has a relatively long, narrow radula, the other group has a shorter, wider radula. The most fundamental differences between these two groups were not apparent until recently. In fact, Coovert (1989b:31) erroneously assigned the genera al- lied to Austroginella to the long, narrow radulate group based solely on radular width. The first major difference noted between these two groups was the discovery of paired, wing-like extensions of the subradular membrane, found only in species with a long, narrow radula (and not in genera allied to Aus- troginella). These structures, which persist in routine tnaceration with KOH, were first noticed by Dean Hew- ish (pers. conmi), and later confirmed by us in a number of species. They were originally called "membranous wings," and appeared to be attached anteriorly to the G. A. Coovert and H. K. Coovert, 1995 Page 53 radular ribbon at the bending plane. They are not readily visible after permanent mounting on microscope slides. We follow Coan (1965) in useage of the term rachidian plate rather than "rachidian teeth" because of the wide, multicusped nature of these structures in marginelliform radulae. We discovered that two different conditions of ra- chidian plate development occur on the posterior end of the radula where plates are initially secreted. Species with a long, narrow radula have a series of at least 6, and up to 12 or more, gradually formed rachidian plates at the posterior end, which are enclosed in a radular sac. These plates, initially narrower and rudimentary, are extremely thin, clear, and very difficult to discern in slide preparations. Progressing anteriorly, these plates become wider and more visible, first developing medially, with lateral portions forming next, finally followed by full cusp development. We refer to these separate undevel- oped plates as nascent rachidian plates, and the gradually developing series as a developmental series. Just anterior to the nascent plates, a series of up to 40 or more fully formed plates are distinctly brown-tinged, followed by fully developed, clear, normal-appearing plates. In con- trast, species with a short, wide radula have only one or two nascent plates posteriorly. These plates are fully formed with completely developed cusps, but are ex- tremely thin, clear, and often very difficult to discern in microscope slide preparations. The first plate often ap- pears as a "ghost image." We term these preformed nascent rachidian plates. Just anterior to these nascent plates, a series of 4 to 5 fully formed brown-tinged plates are followed by fully developed, clear, normal-appearing plates. The buccal pouch, characteristic of marginellids in the strict sense (see Internal Anatomy section), contains the posterior end of the radula in this group. Thus, two fundamentally different conditions exist in both overall structure and development of the posterior end of mar- ginelliform gastropod radulae. In our search for taxonomic characters of marginel- liform radulae, we noted that Ponder (1970:70) reported two completely separate odontophoral cartilages in Gib- berula (as Diluculum sp.). A pair of anteriorly fused odontophoral cartilages (described as a single cartilage in Volvarina taeniolata. see Fretter, 1976:329) were de- scribed from all other studied species. None have been illustrated. We dissected Persicula interruptolineata (Megerle von Miihlfeld, 1816), a close relative of Gib- berula, to confirm the presence of separate odontophoral cartilages. Rather than underlying the radular ribbon as in typical marginellids, the two separate odontophoral cartilages of Persicula each fit into one of the "mem- branous wings" that flank the radular ribbon (figures 15, 16). Further w ork determined the shape and attachments of these wing-like membranous structures. These paired, hood-like extensions of the subradular membrane (more accurately termed the cuticle, see Hyman, 1967:236), which serve to receive the odontophoral cartilages, are here referred to as odontophoral cartilage hoods (figures 13-16). Nothing resembling these structures has been Table 2. .Anatomical charactt-r states utilized in Table S Buccal poucli 0 — absent I — present 2. Neogastropc id radular sac 0 — present 1 — absent 3 General radular t\ pe 0 — cystiscid 1 — marginellid 2 — absent 4 Specific radular type 0 — absent 3 — Type 3 radula 5 — Type 5 radula 6 — Type 6 radula 6m — modified Type 6 radula 7 — Type 7 radula .5 Odontophoral cartilages 0 — separate 1 — anteriorly fused 2 — fused both anteriorly and posteriorly 3 — absent 6. Siphon 0 — attached left of head 1 — fused to head 7 Valve of I.eiblein 0 — present, no bypass tube 1 — present, with bypass tube 2 — absent 8, Esophageal caecum 0 — absent 1 — present 9. Gland ol Leiblein 0 — small, narrow gland, no ter- minal bulb 1 — large, sacculate gland, no terminal bulb 2 — with terminal bulb and long, convoluted duct 10 Duct of gland of Lei- 0 — empties into esophagus pos- blcui terior to nerve ring 1 — passes through nerve ring, emptying into anterior end of proboscis either into buc- cal cavity or anterior end of anterior esophagus 11 Paired salivary glands 0 — ascinous 1 — tubular 12. Paired salivary gland at- 0 — attached to esophagus tachment 1 — free of esophagus 13. Paired salivary glands, 0 — embedded in walls of esoph- ducts agus 1 — attached to esophagus 2 — free of esophagus 14. Accessory salivar\ glands 0 — absent 1 — single gland present reported in marginellids with a short, wide radula. A radula from a specimen of Priinum aff. aletes Roth, 1978, which was not macerated in KOH, revealed anterolateral extensions of the radular ribbon cuticle. These, however, were narrower lateral flanges that differ functionally from the odontophoral cartilage hoods of Persicula. These ex- tensions of the cuticle apparently help hold the radular ribbon in correct alignment with the underlying odon- tophoral cartilage. We term these structures marginal cuticular flanges (figures 17-20). Unhke odontophoral cartilage hoods, they are lost during routine maceration Page 54 THE NAUTILUS, Vol. 109. Nos. 2 & 3 Table 3. Suiiiniar) ol aiiatniiiical icatures of margiiielliform gastropods ('liaractcr stales given in Table 2 Sources of data given in Tabic 4 ''' indicates character state unknown or uncertain. Cvstiscidae Character states 10 11 12 13 14 Pcrsicula inlerruplolineala Pcrsicula masirana Gibberula sp. 0? Marginellidae Margin£llorm gigas 1 7 7 1 0 0 1 0 1 1 ^ 0 Serrala mustclirm 6 m 0 1 0 2 0 1 0 0 1 Austri'ginella johnstoni 5 7 1 0 2 0 1 0 0 0 Auslroginella muscaria 5 7 1 0 2 0 1 0 0 0 Mesoginella pygmaea 5 0 1 0 2 0 1 0 0 0 Volvarma avena 6 2 0 2 2 0 I 1 1 Voh'orina lacniolala 6 2 7 2 2 0 1 2 1 Prunum aides 6 1 0 2 2 1 1 2 0 Prunum gultalum 6 1 0 2 2 0 1 1 0 Prunum marginatum 6 7 1 2 2 0 1 2 0 Prunum marlint 6 7 0 2 2 1 1 2 0 Prunum prunum 6 2 0 2 2 1 1 I 1 Prunum ruosevelli 6 2 0 2 2 1 1 1 0 BuUala buUala 6 7 0 2 2 7 7 7 9 llyalina hyalina 0 2 0 3 0 7 27 17 1 7 7 7 llyalina pallida 0 2 0 3 0 2 2 1? 1 2 1 Douimargo cairetrna 0 2 0 3 0 2 0 2 I 1 2 1 DciUimargo eburneola 0 2 0 3 0 2 0 2 0 1 2 1 Marginclla desjardini 0 2 0 3 0 2 0 2 7 1 9 1 Marginella glabella 0 2 0 3 0 2 0 2 1 1 2 1 Marginella sehasliani 0 2 n 3 0 2 0 2 1 1 2 1 G. A. Coovert and H. K. Coovert, 1995 Page 55 odontophoral cartilage hood marginal cuticular flange 14 odontophoral cartilage Figures 13-16. Persicula interruptolineata (Megerle von Miihlfeld, 1816), showing details of cystiscid radular morphology. GAC 1581, data in Table 1, Rachidian width enlarged ISC^c for clarity. 13. Left lateral view, odontophoral cartilages removed, only posterior end of radular sheath shown. 14. Same, dorsal view. 15. Dorsal view, odontophoral cartilages in place, muscles and connective tissue removed. 16. Same, anterior view. Figures 17-20. Pntnum aff. aletes Roth, 1978, showing details of marginellid radular morphology. GAC 1518, data in Table 1. 17. Left lateral view, odontophoral cartilage removed. 18. Same, dorsal view. 19. Dorsal view, odontophoral cartilage in place, muscles and connective tissue removed. 20. Same, anterior view. in KOH, suggesting additional differences in composi- tion. Morphological evidence suggests that there are two fundamentally different types of radulae. The first type has a very long, triserial or uniserial, C-shaped radula, composed of 80 to 200 or more very narrow, usually overlapping, arched rachidian plates, with relatively few (3 to 15) cusps. This type of radula has a pair of odon- tophoral cartilage hoods, each receiving a separate odon- tophoral cartilage, which flank the radular ribbon an- teriorly. The posterior end of the radula begins as a developmental series of nascent plates and is enclosed in a typical neogastropod radular sac. This type of radula is found in all species here assigned to the Cystiscidae and is termed a cystiscid radula (figures 13-16). The second type has a uniserial, J-shaped ribbon composed of 10 to 80 broad, overlapping, flat rachidian plates, usually with numerous cusps (9 to 80, rarely 4 to 7). This type of radula has marginal cuticular flanges, and the odontophoral cartilages are fused anteriorly into a single structure underlying the radular ribbon. The posterior end of this type of radula starts with 1 or 2 preformed nascent rachidian plates and is enclosed in a marginellid buccal pouch. This second type is found in all radulate species here assigned to the Marginellidae s. str., and is termed a marginellid radula (figures 17-20). Coovert (1989b:31-33) divided marginelliform radu- lae into seven distinct types based on characters of the rachidian plates and the presence or absence of lateral teeth. In addition to these seven types, which are dis- cussed below, two new types are recognized. The ori- entation of rachidian plates refers to the large, dorsal Page 56 THE NAUTILUS. Vol. 109, Nos. 2 & 3 Marginellid Radulae Type 5 Modified Type 6 Cystiscid Radulae Type 1 Type 2 Type 3 Figures 21-24. Cystiscid radular types 21. Pleniocystisciis jeivettii (Carpenter, 1857). example of Type 1 radula G,'\C radula no 093, Pirates Cove, San Luis Obispo, California, intertidal. Ex. M1823, shell length 4.9 mm. 22. Cystiscus angasi (Crosse, 1870), example of Type 2 radula. Hewish radula U, Lonsdale Bight, Victoria, Australia. Ex. NMV F53748, shell length 1.7 mm. 23. Pcrsicula persicula (Linne, 1758), example of Type 3 radula. GAC radula no, 183, Niodior, Senegal. Ex. AMNH 173759, shell length 19.8 mm 24. Granulina hadria (Dall, 1889), example of Type -4 radula. GAC radula no. 038, Tampa Bay, Pinellas Co., Florida, sand and mud. Ex GAC Ace # 27-82, shell length 2.2 mm. Figures 25-30. Marginellid radular types 25. Atistroginella muscaria (Lamarck, 1822), example of Type 5 radula GAC radula no. 181, Lakes Entrance, Victoria, Australia. Ex, DMNH 12435, shell length 13 3 mm. 26. Serrata Iranslata (Redfield, 1870), example of Modified Type 6 radula GAC radula no 304. Raiatea, French Polynesia. Ex. GAC Ace, # 2-91, shell length 4.3 mm. 27. Prunum apicinum (Menke, 1828), example of Type 6 radula, GAC radula no. 147, Sunshine Key, Florida Keys, Monroe Co,, Florida, Ex, Peggy Williams colln,, shell length 95 mm. 28. Margincllona gigas (Martens, 1904), example of Type 7 radula. After Thiele (1904;pl, 9, fig, 64), Ex, Holotype, shell length 80 — 100 mm. 29. Serrataginclla spryi (Clover, 1974), example of Type 8 radula, G.f^C radula no, 305, off Nacala Bay, N, Mozambique, dredged in 20-30 m, sand, Ex. GAC; Ace. # 6-93, shell length 8,0 mm 30. Hydroginella tridenlata (Tate, 1878), example of Type 9 radula G.AC radula no 306, Margaret River, Western Australia, in sand, 4 6 m Ex G.^C \cc. # 13-90, shell length 7,8 mm. section of the ribbon, not the veiitrally folded anterior end. This orientation is such that the cutting edge of each rachidian plate is posterior, and the basal edge, where the plate is attached to the membrane, is anterior. Type 1 through Type 4 radulae are all c\ sticid radulae, possessing the characteristics previously defined. Type 1 ("Triseriai Type") radulae (figure 21 ) are triserial, with simple, thin, weak, lateral teeth that narrow to a single, posteriorly-pointed cusp. The long, narrow radula is composed of 66 to 103 rows of overlapping rachidian plates. These plates are very narrow, weakly arched, with 3 to 7 sharp cusps along cutting edge. The basal edge of each rachidian plate is slightly concave, resulting in a weakly Ll-shaped plate, Odontophoral cartilage hoods were noted by us as well as by Dean Hewish (pers, comm, ), but the nature of the odontophoral cartilages is unknown. The genus previously referred to as "Cystiscus" (Coo- vert, 1989b), described herein as new, is the only one to possess a Type 1 radula (radulae of 3 species studied). Type 2 radulae (figure 22) are uniserial, consisting of a G. A. Coovert and H. K. Coovert, 1995 Page 57 long, narrow series of over 80 to 220 rachidian plates. These plates are overlapping (rarely separate), very nar- row, weakly to strongly arched, with 4 to 15 very strong cusps along cutting edge. The basal edge of each plate is slightly concave, nearly straight, or convex medially, resulting in a crescent-shaped, subrectanguiar, or asym- metrically V-shaped plate. Odontophoral cartilage hoods were noted in many species of Cystisctis. but were un- known and unobserved at the time Crithe radulae were extracted. The nature of the odontophoral cartilages is unknown. Cystiscus (radulae of 14 species studied) and Crithe (radulae of 1 species studied) both have a Type 2 ("Crithe / Cystiscus Type") radula. The radula of Extra is unknown, but presumed to be similar. Type 3 radulae (figure 23) are uniserial, consisting of a long, narrow series of over 80 to 209 rachidian plates. These moderately to strongly overlapping plates are nar- row, moderately to strongh arched, with 5 to 14 sharp cusps along cutting edge. The central cusp is often the strongest. The basal edge of each plate is strongly con- cave, resulting in a U- or V-shaped plate. The reports of wishbone-shaped rachidian plates (Barnard, 1962:14-15; Ponder, 1970:70; Ponder, 1973:fig. 2) are inaccurate and due to an artifact of observation (see discussion in Coo- vert, 1989b: 12-13). Odontophoral cartilage hoods were noted in all three included genera, with a pair of separate odontophoral cartilages noted in Persicula and Gibber- ula. Persicula (radulae of 14 species studied), Gibberula (radulae of 21 species studied), and Canalispira (radulae of 2 species studied) all have a Type 3 ("Persicula Type") radula. Type 4 radulae (figure 24) are uniserial, consisting of a long, narrow series of at least 90 to 159 rachidian plates. These usually completely non-overlapping plates are nar- row, moderately arched, with 9 to 12 clustered, raised denticles located in staggered positions on the dorsal sur- face, with a single (or asymmetrically paired) strong central cusp protruding on the cutting edge. The basal edge of each plate is weakly concave, resulting in a weakly V-shaped plate. The rachidian plates are asym- metrical and alternate as mirror images (figure 24). Al- though odontophoral cartilage hoods have not been ob- served in representatives of this group, their radulae are otherwise typically cystiscid. A broad, membranous flange on either side and extending the whole length of the radula, effectively doubling the width, has been observed (Dean Hewish, pers. comm.) when using the enzyme Pronase for maceration. This is a less destructive method than the use of KOH. This flange is certainly homologous to odontophoral cartilage hoods, but the nature of the odontophoral cartilages is unknown. No further anatom- ical information is available. Granulina (radulae of 12 species studied) and Fugnu.s (radulae of 2 species studied) both have a Type 4 ("Granulina Type") radula. Type 5 through Type 9 radulae are all marginellid rad- ulae, possessing the characteristics as previously defined. Type 5 radulae (figure 25) are uniserial, consisting of a relatively short, broad ribbon of 19 to 75 rachidian plates. These usually overlapping plates are moderately wide, weakly arched, with 8 to 22 strong cusps along cutting edge. The central cusp is often the strongest. There are often pits on adjacent plates to receive the tips of some of the cusps, especially the central. The basal edge of each rachidian plate is generally slightly concave, re- sulting in a rectangular to chevron-shaped plate. The odontophoral cartilages in Mesoginella and Austrogi- nella were described as fused anteriorly (Ponder, 1970: 62; Ponder & Taylor, 1992:320), but no detailed data exists as to marginal cuticular flanges. Protoginella (rad- ula of 1 species studied), Alaginella (radulae of 9 species studied), Austroginclla (radulae of 5 species studied), Mesoginella (radulae of 16 species studied), Ovaginella (radulae of 2 species studied), and Balanetta (radula of 1 species studied) all have a Type 5 ("Austroginclla / Mesoginella Type") radula. The radula of Closia is un- know n, but presumed to be similar. Type 6 radulae (figure 27) are uniserial, consisting of a relatively short, broad ribbon of 24 to 75 overlapping rachidian plates. These plates are broad, nearly flat, with 9 to 45 (exceptionally 5) sharp cusps along cutting edge. There are often pits on adjacent plates to receive the tips of some or all of the cusps. The basal edge of each plate is generally straight, resulting in an elongate, rectangular "comb-like" plate. The odontophoral cartilages are fused anteriorly. They are fused posteriorly in most species (but not in Prunum aff. aletes and P. guttatum), leaving a narrow, medial, longitudinal posterior slit in most Prun- um and Volvarina (figures 38-42). (See following section on internal anatomy.) A single cartilage was described for Volvarina taeniolata (cf. Fretter, 1976:329). The marginal cuticular flanges of Prunum aff. aletes, de- scribed above, are very similar in other species of Prun- um we examined. Prunum (radulae of 23 species stud- ied), Volvarina (radulae of 42 species studied), Rivo- marginella (radulae of 2 species studied), Bullata (radula of 1 species studied), and Cryptospira (radulae of 4 spe- cies studied) all have a Type 6 ("Prunum / Volvarina Type") radula. Most of these are discussed and figured in Coovert and Coovert (1990). Modified Type 6 radulae (figure 26) are similar to typical Type 6, but differ in having a shorter ribbon composed of only 13 to 35 plates, averaging more cusps (22 to 59) that project from a somewhat sinuous posterior edge, and in often having thinner and more fragile rachidian plates. Pits on adjacent plates to receive cusp tips were not observed as in many typical Type 6 radulae. The odon- tophoral cartilages of Serrata (as Haloginella) were de- scribed by Ponder (1970:66) as fused except for a very short posterior portion. Marginal cuticular flanges are unknown. Serrata (radulae of 8 species studied) is the only genus to have a Modified Type 6 ("Serrata Type ) radula. Type 7 radulae (figure 28) are uniserial, consisting of a relatively short, very broad ribbon of 56 to 80 overlap- ping rachidian plates. These plates are very broad, nearly Page 58 THE NAUTILUS, Vol. 109, Nos. 2 & 3 flat, with numerous (58 to 85) sharp cusps along cutting edge. The basal edge of each plate is generally straight, resulting in an elongate, rectangular "comb-like" plate. Odontophoral cartilages and marginal cuticular flanges are unknown. The monotypic genera Marginellona and Afrivuluta both have a Type 7 {" Afrivoluta Type") rad- ula. Type 8 radulae (figure 29), here described, are uniserial, consisting of a relatively short, broad ribbon of 38 over- lapping rachidian plates. These plates are broad, nearly flat, with 9 to 10 sharp cusps on the cutting edge. The entire posterior edge, including all edges of the main cusps, with numerous (total of ca. 70) small, secondary cusps, giving the main cusps a serrated appearance. Pits for reception of adjacent cusp tips were not observed. The basal edge of each plate is slightly indented medially, resulting in a very shallowly V-shaped plate. Odonto- phoral cartilages and marginal cuticular flanges are un- known. The Type 8 ("Serrataginella Type") radula is unlike any others known, and is represented by "Mar- ginella" spryi Clover, 1974, type species of a monotypic genus herein described as new. Type 9 radulae (figure 30), first described by Bouchet (1989:79, fig. 3), are uniserial, consisting of a very short, narrow, greatly reduced ribbon of 10 to 30 rachidian plates. These weak plates overlap and are nearly flat, with 4 to 7 cusps on the cutting edge. Pits for reception of adjacent cusp tips are absent. The basal edge of each plate is straight, resulting in a subquadrate plate. Odon- tophoral cartilages and marginal cuticular flanges are unknown. Hydroginella (radulae of 4 species studied) is the only genus to have a Type 9 ("Hydroginella Type") radula. One species is known to be parasitically associated with fishes (Bouchet, 1989). Judging from the very sim- ilar radulae of other species, it seems likely that all may have such an association. A third major group, comprised of non-radulate spe- cies, completely lacks a buccal mass, odontophore, and radula. This was confirmed by dissections of species in the following genera: Dentimargo (Ponder, 1970:67, plus species reported herein), Marginella s. str. (Graham, 1966: 139, Coan & Roth, 1976:220, plus species reported here- in). Glabella (Coan & Roth, 1976:220), and Hyalina (Coan & Roth, 1976:220, and herein). Other species have been subsequently reported to lack a radula, some based on unsuccessful radular extraction with KOH (see Coovert, 1989b tor review). Thus, a total of 31 species have been inferred or demonstrated to lack a radula. We have at- tempted routine radular extraction on separate occasions from two specimens of the type species of Eratoidea, E. margarita (Kiener, 1834), and have not found a radula. Although not definitive, it is suggested tiiat this genus is also non-radulate. This assemblage of non-radulate gen- era is an artificial group, with Hyalina l)ei()Mging to a different lineage (see discussion in systematic section). Internal Anatomy: Prior to our studies, only 13 species of marginelliform gastropods had been studied anatom- ically. Harasewych and Kantor (1991:tables 2, 3) sum- marized anatomical data of the 11 species known at that time. Ponder and Taylor (1992) subsequently presented data for two species of Austroginella. Few taxonomic conclusions could be drawn from such a relatively small sample because the anatomy varied so greatly. Our dis- sections added anatomical data for 11 additional species, and further data for one previously reported species (Ta- bles 1-4). Combined with conchological and radular data, this greatly expanded body of anatomical information enabled taxonomic groupings to become evident, making a reliable and stable classification possible. Details of the foregut were emphasized because of their greater taxo- nomic value. Our primary objective is a presentation of the higher classification, therefore, more detailed reports on the anatomy of individual species will be published elsewhere. A discussion of foregut anatomy follows, along with a summary of known anatomical information for these 24 species. The proboscis of marginelliform gastropods is rela- tively short and pleurembolic. The retractor muscles are inserted on the sides of the proboscis, resulting in the basal portion being invaginated, forming a proboscis sac (Fretter & Graham, 1962:150). The proboscis varies somewhat in shape among genera. The small mouth is always situated terminally at the anterior end. Most gen- era have a proboscis that is rounded or blunt distally, and, when retracted, is generally about two-and-one-half to four times as long as broad. When feeding, the pro- boscis may be extended to a length approximating the length of the shell in at least some species (Fretter, 1976: 329). Both species of Dentimargo studied have a distally pointed proboscis about four times as long as broad when retracted (figure 33). Serrata aff. mustelina has a more bulbous, distally rounded proboscis about one-and-a-half times as long as broad. Gibberula and Persicula (figures 35, 79) both have a short, broad proboscis about two times as long as broad. The mouth opens directly into the buccal cavity. No jaws have been observed in any marginelliform species. In radulate species, the buccal cavity leads posterodor- sally to the anterior esophagus and posteroventrally to the buccal mass. In non-radulate species, the buccal cav- ity leads with little or no differentiation to the anterior esophagus. The most significant hallmark of the family Margi- nellidae, in the strict sense, is the possession of a "buccal pouch" or "buccal caecum" in radulate species. Both Fretter (1976:333) and Harasewych and Kantor (1991: 16) recognized the significance of this structure and its presence only in marginellids and toxoglossans. The buc- cal pouch (figure 31), the term we prefer for this struc- ture, was first discovered and named by Graham (1966: 137), who noted that "in marginellids the most interest- ing feature of the gut is the situation of the buccal mass, wholly withdrawn into a caecum from its typical place on the floor of the buccal cavity (op. cit., p. 147)." The l)uccal mass, described as "rather reduced" by Graham (1966:137) and consisting of the odontofihoral cartilages. G. A. Coovert and H. K. Coovert, 1995 Page 59 Table t. Sources of anatomical data utilizcil in Tabic 3 Species (current nonicuclaturcl Source Cystiscidae Pcnicula intcrruptolincata (Megcrle von Miihlleld, 1816) Pcrsiciila masirana Roth 6i Petit, 1972 Cibherula sp Marginellidae Marginvllona gigaa (Martens, 1904) Serrata aff mustclina (Angas, 1871) Amtroginclla johnstoni (Petterd, 1884) Austroginclla muscaria (Lamarck, 1822) Mesoginrlla pygmaca (G. B. SouerhN II, 184(i) Volvarind all ciicnci (Kiener, 1834) Volvarina tacmulata Morcli, ISHO Priinnm aff. alctcs Rotli, 1978 Pruntim guttalum (Dill\v\ii. 1817) Pruntim marginatum (Born, 1778) Pniniim martini (Petit, 18.53) Pninum prunum (Gmelin, 1791) Pruniuii rooscvciti (Bartscli & Relider, 1939) Bullala huUata (Born, 1778) Htialina lujalina (Tfiiele, 1913) Hi/alina pallida (Linne, 17.58) Dcntimargu cairoma (Brookes, 1924) Dcntimargo cburneola (Conrad, 1834) Marginclla dcsjardini Marche-Marehad, 1957 Marginella glabella (Linne, 1758) Marginclla scliantiani Marclie-Marcliad & Rosso, 1979 herein; see Table 1 for data herein; see Table 1 for data Ponder (1970), as Diluculuni sp, Harasewych and Kantor (1991) Ponder (1970), as Volvarina (llalogindia) mustelina, the New Zealand species differs from the Australian species; see Coovert (1989b: 16) for comments Ponder and Tavlor (1992) Ponder and Taylor (1992) Ponder (1970), as Mesoginella iSinuginclla) pygmaea herein; see Table 1 for data Fretter (1976) herein; see Table 1 for data herein; see Table 1 for data Graham (1966), as Marginella mariginata Marcus and Marcus (1968), as Mariginella fratercultis herein; see Table 1 for data herein; see Table 1 lor data herein; see Table 1 for tlata Eales (1923), as Marignella hyalina Coan and Roth (1976) jihis herein; see Table 1 for data Ponder (1970), as \'olvarinella cairoma herein; see Table 1 for data Graham (1966) herein; see Table 1 for data herein; see Table 1 for data radula, and associated connective tissue and muscles, is contained in the buccal pouch. What characterizes the buccal pouch is the sphincter at its anterior opening into the buccal cavity and the fact that it is a separate caecum or sac that is blind posteriorly, where the radula is ini- tially secreted. This is further discussed by Graham (1966; 138), who noted that radular formation occurs "at the innermost end of the pouch, which therefore corresponds to the radular sac of more normal gastropods," Ponder (1970:76) also noted that the buccal pouch is derived from the radular sac. Fretter (1976:329) observed that the "teeth arise at the innermost end of the pouch where there is no clear demarcation of the radular sac typical of prosobranchs." The blind end of a buccal pouch thus corresponds to the posterior end of the radular sac. Our observations of Prunum, Volvarina, and Bullata species confirmed the presence of a buccal pouch. The genera Serrata. Mesoginella, and Austroginclla, dissected by Ponder (1970) and Ponder and Taylor (1992), likewise possess a true marginellid buccal pouch. In the true mar- ginellids, the odontophore extends beyond the posterior end of the radula. The absence of a buccal pouch in non-radulate species has been confirmed by dissections of species of Hyalina, Dentimargo, and Marginella (figures 32-34). Graham (1966:139) described what he believed to be a vestigial buccal pouch in Marginella desjardini. Ponder (1970: 76) considered this to be the accessory salivary gland. Our dissections of Marginella glabella (figure 34) and M. sebastiani, both close relatives of M. desjardini, con- firmed this contention, because an accessory salivary gland was found in both species, but neither had a vestigial buccal pouch. Ponder's (1970: fig. 3B) dissection of Den- timargo cairoma, and our dissection of D. eburneola (figure 33), show the complete absence of a buccal pouch. There is insufficient information about Hyalina hyalina in Eales (1923) to draw any conclusions. Coan and Roth (1976:220) likewise did not provide sufficient detail for H. pallida, although they did mention the absence of a radula and odontophore. Our dissection of H. pallida (figure 32), confirms the absence of a buccal pouch. A typical neogastropod radular sac enclosing the pos- terior end of the radula was found in both species of Persicula we dissected (figure .35). In Persicula inter- ruptolineata and in the Gibberula studied by Ponder (1970:fig. E, r.s), the radular sac is readily distinguished because it protrudes from the posterior end of the pro- boscis in a characteristic downward curve (figures 35, 79). It is further characterized by a slightly bulbous pos- terior end which encases the nascent end of the radular ribbon (cjv. previous section). The radular sac extends well behind the odontophore and is an extension of the radular sheath, formed from the subradular membrane. The buccal mass in Persicida masirana is morphologi- Page 60 THE NAUTILUS, Vol. 109, Nos. 2 & 3 gland of Leiblein esophageal caecum esophagus duct of gland of Leiblein mouth buccal pou paired salivary glands buccal mass 31 gland of Leiblein esophagus mouth duct of gland of Leiblein paired salivary glands accessory salivary gland Figures 31-32. Anatomical features dI llie foregiit of Pruiiini Data in Tahle I 31. Pninum aff. aleies Rotli, 1978 12-4 X. 32. Hyalinu pallida (Linne, 1758). 15.3 .\, cally the same, but the proboscis is simply longer, com- pletely containing the radular sac. The characteristic shape and protrusion of the radular sac seems to be fairly typical for most Neogastropods (including Olividae), with the prominent exception being the Marginellidae, s. str. The radular sac thus differs in several respects from the buccal pouch of iiiargiiiellids, especially in the lack of an anterior sphincter. All cystiscid genera studied have the same general type of radula, previously termed a cystiscid radula ((jv.). Based on the similar na.sceiit ends of their radulae, all cystiscid genera are a.ssumed to pos- sess a true radular sac similar to that found in Persictila and Gibbcrula. A radular sac has actually been observed in several incompletely cleared cystiscid radulae, in- cluding the triserial group "Cystiscus," named as a new genus herein Odontophorai cartilages, which we prefer over "bol- sters, " "odontophore, or "cartilages," is a more specific term and clearly differentiates these structures from the odontophore, which is often used as a general term for the entire radular mass. These cartilages support and stiffen the radular mass and are composed of large Ley- G. A. Coovert and H. K. Coovert, 1995 Page 61 esophagus nerve ring gland of Leiblein mouth accessory salivary gland duct of gland of Leiblein pairea salivary gland mouth esophagus accessory salivary gland duct of gland of Leiblein gland of Leiblein paired salivary glands 34 mouth valve of Leiblein esophagus buccal mass accessory salivary gland gland of Leiblein 35 Figures 33-35. Anatomical features of the foregut of Marginellini and Persiculinae^ Data in Table 1 33. Dentimargo ebiirneola (Conrad, 1834). 22.4 X. 34. Marginella glabella (Linne, 1758). 5.2 X. 35. Persicula interruptolineata (Megerle von Miihlfeld, 1816). 16.8 X. Page 62 THE NAUTILUS, Vol. 109, Nos. 2 & 3 39 42 Figures 36-42. Odontophoral cartilages of cystiscids and marginellids Data in Table 1 36. Persicula interruptolineata. GAC 1581A. Length 0 97 mm 37. Persicula masirana. GAC Ace. # 4-94B Length LOl mm 38. Volvarina aff. avena. GAC 1172B Length 0 88 mm. 39. Prtuwm aff. aides. GAC 1518B. Length L50 mm. 40. Prunum guttatum. GAC 1172A. Length 0.75 mm. 4L Prunum prunum. GAC 1582. Length 2.12 mm. 42. Prunum roosevelti. GAC Ace # 26-88A. Length 1.67 mm. dig cells (chondroid tissue) by which they are easily rec- ognized. Two fundamentally different conditions also exist in this character complex. In all true marginellids examined, the two odonto- phoral cartilages are fused at least anteriorly, effectively functioning as a single structure underlying the radula. Ponder (1970:62) described the odontophoral cartilages in Mesoginella pijgmaea as "fused into a single structure in the front half of the odontophore but posteriorly are connected by a transverse muscle in the usual way." Ponder (1970:66) described similarly fused cartilages in Serrata aff. mustelina, which are separated for only a short distance posteriorly. The two species of Austrogi- nella discussed by Ponder and Taylor (1992:320) are likewise described as having anteriorly fused cartilages. The odontophoral cartilages in Marginellona were not described by Harasewych and Kantor (1991). Fretter (1976:329) simply described a single cartilage for Vol- varina taeniolata. Our dissection of V. aff. avena re- vealed a pair of cartilages fused both anteriorly and posteriorly (figure 38). They are slightly bowed medially, leaving a medial slit about half the length of the entire structure, with the anterior margins raised and thick- ened, and truncated anteriorly. We suspect that V. tae- niolata may be similarly structured. The closely related genus Prunum shows a similar condition. These struc- tures were insufficiently described in Graham (1966) and Marcus and Marcus (1968) to draw any conclusions. Our dissections of four species of Prunum revealed that they all have strongly raised, thickened anterolateral margins, effectively forming a trough or channel for the radular ribbon (figure 20). The odontophoral cartilages in P. roosevelti (figure 42) are relatively narrow, rounded an- teriorly, completely fused on the very long and narrow anterior half, and slightK bowed and narrowly fused posteriorly, confining a medial slit to the posterior half. The odontophoral cartilages in Prunum prunum (figure 41) are relatively short and broad, fused anteriorK and posteriorly, with a narrow medial slit. The anterior end is shallowly emarginate. Prunum guttatum (figure 40) and P. aff. aletes (figure 39) differ because their odon- tophoral cartilages are \'-shaped, and narrowly fused anteriorly, but not posteriorly. The fused cartilages in P. guttatum (figure 40) ha\e the posterior tips slightly con- vergent but not touching, with the anterior end truncate. In P. aff. aletes (figure 39), the anterior end is truncate to shallowly emarginate, and the posterior tips are di- vergent, forming a distinct \'-shape, but in situ (figure 19) connective and muscle tissue hold these posterior tips closer together. In all cases, this fused pair of odonto- phoral cartilages underlies the main part of the radula (figures 19, 20). The radula wraps around the anterior G. A. Coovert and H. K. Coovert, 1995 Page 63 end of the odontophore, tliis point being effectively termed the bending plane. The shorter anterior portion of the radular ribbon, often showing broken cusps, thus hes under the anterior end of the odontophore, giving it the characteristic J-shape. Our data indicates that the odontophoral cartilage shape is consistent within each species, but differs between them, and may prove to be a useful taxonomic character. All cystiscid genera examined have two completely separate odontophoral cartilages. Each cartilage fits into one of the lateral odontophoral cartilage hoods of the subradular membrane, flanking and not underlying the radula (figures 15, 16). Ponder (1970:70) described the odontophoral cartilages and muscle attachments for a species of Gibberula. Our dissections of Persicula inter- ruptolineata (figure 36) revealed two separate subtrian- gular cartilages that are broad and rounded anteriorly, and narrowed posteriorly. In P. masirana (figure 37), the separate cartilages are elongate, rather uniform in width throughout, and pointed anteriorly. The radulae of near- ly all other cystiscid genera have been described earlier in this paper, and most have odontophoral cartilage hoods. On this basis, we infer that they possess two separate odontophoral cartilages. The anterior esophagus in radulate genera is situated dorsally within the proboscis, the buccal mass lying below it. In non-radulate species, the anterior esophagus is more centrally placed. Upon exiting the retracted proboscis posteriorly, the esophagus generally makes a sharp S-bend prior to passing through the nerve ring. A valve of Lei- blein, when present, is located anterior to the circum- esophageal nerve ring. Ponder and Taylor (1992:321) described the valve of Leiblein in Austroginella as being partially encased in the nerve ring anteriorly (figure 80). Serrata and Mesoginella have a well-developed valve located just anterior to the nerve ring. These three genera have a narrow, non-glandular bypass tube that is adher- ent ventrally to the valve of Leiblein. Marginellona has a large valve of Leiblein situated anterior to the nerve ring (figure 80). The two Persicula species we dissected also have a distinct valve of Leiblein, placed well anterior to the nerve ring (figure 35). No bypass tube was found. Gibberula has a similarly placed valve of Leiblein, but has a separate glandular tube that bypasses the valve (figure 79). Volvarina, Hyalina, Pninuvi, Btillata. Den- tiniargo, and Marginella all lack a valve of Leiblein. Marcus and Marcus (1968:65) discussed an "individually inconstant valve of Leiblein" in Prunum martini. In some Prunum species we dissected, what at first ap- peared to be a possible valve of Leiblein simply turned out to be food lumps in the esophagus that were incon- sistent in their location or occurrence. We believe that the so-called "individually inconstant valve of Leiblein ' in Prunum martini were simply food lumps in the esoph- agus of some individuals. The esophagus narrows upon entering the nerve ring anteriorly, resuming its original size after exiting pos- teriorly. A much enlarged sac-like esophageal caecum was found just posterior to the nerve ring in all species of Volvarina. Prunum (figure 31), and Bullata studied. This structure was first described in Prunum margina- tum as an "oesophageal caecum" (Graham, 1966:135, fig. 1). Eales (1923) did not mention this structure in Hyalina hyalina, nor was it mentioned by Coan and Roth (1976) for H. pallida. Our dissection of H. pallida (figure 32) revealed a thin-walled, but very distinct, esophageal caecum. An esophageal caecum is absent in all other marginelliform genera that have been studied anatom- ically. The most conspicuous foregut gland is the gland of Leiblein, also called the "unpaired gland, "unpaired foregut gland, or "poison gland." This gland occurs in three different forms. In Persicula (figure 35) and Gib- berula (figure 79), this gland is a small, short, often slight- ly folded gland emptying directly into the posterior end of the mid-esophagus, well posterior to the nerve ring. In Marginellona (figure 80), the gland of Leiblein is extremely large, sacculate, broad anteriorly and tapering posteriorly (Harasewych and Kantor, 1991:11). It also empties directly into the mid-esophagus posterior to the nerve ring. All the remaining marginelliform genera studied anatomically have a gland of Leiblein with a long, convoluted duct. The gland in these genera ends distally in a terminal bulb that is either rounded or weak- ly to distinctly pointed distally. The terminal bulb is narrow, elongate, and rounded distally in Der^timargo (figure 33). In Marginella, it is shorter, more swollen, and ends in an acuminate, pointed tip (figure 34). In Serrata. Mesoginella, and Austroginella (figure 80), the relatively large duct from the gland of Leiblein, which is about the same diameter as the esophagus, empties into the posterior end of the mid-esophagus just posterior to the nerve ring but does not pass through the nerve ring. These three genera also possess a well-developed valve of Leiblein with a ventral by-pass tube. In Prunum (figure 31), Bullata, Volvarina, Hyalina (figure 32), Den- timargo (figure 33), and Marginella (figure 34), the long, convoluted duct from the gland of Leiblein narrows as it passes through the nerve ring, then continues as a very narrow tube to the anterior end of the proboscis. These six genera all lack a valve of Leiblein. The duct from the gland of Leiblein empties either into the buccal cav- ity or into the extreme anterior end of the anterior esoph- agus, which is usually in very close proximity to the point where the ducts of the paired salivary glands empty into the buccal cavity. Ponder (1970:77, 79) described this duct in Marginella desjardini as opening "ventrally into the oesophagus a little in front of the nerve ring." This is in error because Graham (1966:139) clearly described this duct as opening "to the gut at almost the same level as that at which the salivary ducts reach the same po- sition," which is at the "extreme anterior end of the proboscis, where they discharge, ... into the buccal cavity. ' This is mentioned because apparently a figure in Ponder (1973:fig. 3N) is based on this misconception. This does not weaken Ponder's hypothesis for the formation of the poison gland (Ponder, 1970:77-80), but it indicates that this intermediate stage is presently unknown. Page 64 THE NAUTILUS, Vol. 109. Nos. 2 & 3 All inarginelliform genera have either tubular or as- cinous paired salivary glands associated with the esoph- agus. In many cases, tubular and ascinous salivary glands appear to be found in different species of the same genus, and thus may be of limited taxonomic value in these genera. In all cases, the paired glands send fine, narrow ducts to the anterior end of the proboscis, where they empty into the buccal cavity. The placement of the glands and the attachment of ducts differ between species. In Marginellona. the paired salivary glands are found dor- sal to the gland of Leiblein (figure 80), their ducts run- ning anteriorly along the floor of the proboscis, free of the esophagus. Serrata, Mesoginella, and Austroginella all have glands that are attached to tfie esophagus just anterior to the valve of Leiblein, the ducts embedded in the esophageal walls for their entire length. Volvarina taeniolata has the glands free in the body cavity and the ducts free of the esophagus. In V. aff. avena. the paired salivary glands are relatively large and fully contained within the retracted proboscis, the ducts attached ante- riorly to the esophagus. In Prunum. the paired salivary glands are free of the esophagus. In P. prunum and P. guttatum, the ducts are attached to the walls of the esophagus. In the other Prunum species studied, the ducts lay ventrally along the floor of the proboscis, free of the esophagus. Details on the salivary glands of Bullata bul- lata are unknown. The ducts in H. pallida are free of the esophagus, the glands apparently tubular but poorly preserved in the specimen studied. Both Dentimargo species have the glands largely contained within the pro- boscis. They are presumed to be tubular in D. cairoma. but are ascinous in D. eburneola. Oddly, these glands are not placed laterally in D. eburneola as is usual, but instead are placed dorsally and ventrally, one gland sit- uated above the esophagus, the other below the accessory salivary gland (figure 33). In both species, the glands and their ducts lay free. The paired salivary glands of the three species of Marginella are located outside the pro- boscis near the nerve ring, free from the esophagus. In M. glabella, the ducts are attached to the duct of the gland of Leiblein (figure 34), unlike those of M. sebas- tiani. The paired salivary glands in Persicula and Gib- berula are attached to the esophagus just anterior to the valve of Leiblein (figure 35). The ducts of Persicula are attached to the outside walls of the esophagus, but in Gibberula this is unknown. Ponders illustration (1970: fig. 3K) seems to indicate that these ducts are embedded. Information on these glands anil their ducts is summa- rized in Table 3. The acces.sory salivary gland, when present, is always single and empties into the buccal cavity. It is present in all cystiscids studied, being ascinous in both species of Persicula, but tubular in Gibberula. Within margi- nellids, a tubular accessory salivary gland is present in Serrata, but absent in Austroginella and Mesoginella. A large, ascinous gland is present in Volvarina aff. avena, but appears to l)e tubular in V. taeniolata as described by Fretter (1976:329). Both species of Hijalina possess an accessory salivary gland, which is tuliular in H. hy- alina and apparently so in H. pallida (unreported in Coan & Roth, 1976 and poorly preserved in our specimen). An accessory salivary gland is present in only one species of Prunum, being tubular in P. prunum. The gland is pres- ent and tubular in Dentimargo and Marginella, but ab- sent in Marginellona. Further work, including histology, is needed. See Table 3 for a summary of the occurrence of this gland. Our primary objective was to discern useful taxonomic characters, and we feel that the most useful features are those of the foregut. Thus, the remainder of the digestive system was not studied in any greater detail by us, nor was the reproductive system. A brief summary of pub- lished information on these other systems follows. The presence or absence of an anal gland should be further investigated. This structure was reported as pres- ent and extremely large in Marginellona gigas by Har- asewych and Kantor (1991:11). Serrata aff. mustclina, Mesoginella pygmaea, Dentimargo cairoma (cf. Ponder, 1970), Austroginella johnstoni, A. muscaria (cf. Ponder & Taylor, 1992:322), and Volvarina taeniolata (cf. Fret- ter, 1976:329) were all reported as having an anal gland. The only marginelliform reported to lack an anal gland is the species of Gibberula reported by Ponder (1970: 71). Published accounts of the remaining species did not report the presence or absence of this structure. Main features of the reproductive system were sum- marized in Harasewych and Kantor (1991:tables 2, 3). Published information on the reproductive system is in- complete for a number of species described anatomically. Probably of most interest in the female reproductive system is the presence of three glandular structures or seminal receptacles joining the pallial oviduct between the albumen gland and capsule gland in Dentimargo cairoma (cf. Ponder, 1970:74), whereas only one is pres- ent in Gibberula (op. cit., p. 75). Two such structures are the usual number, as recorded in Mesoginella pygmaea and Serrata aff. mustelina by Ponder (1970:74), Mar- ginellona gigas by Harasewych and Kantor (1991:11), Marginella desjardini by Graham (1966:140), and Vol- varina taeniolata by Fretter (1976:fig. 2A). Information is lacking for the other species. In the male reproductive system, the penis is normally simple, lacking appendages or lobes. But Ponder (1970:72, fig. 4C) described a bi- lobed penis for Dentimargo cairoma that has a sharply pointed posteroapical appendage, which he termed a sheath. Our dissection of D. elmrneola likewise revealed a sharply pointed apical appendage, but the penis was not bilobed. The penis in Bullata Imllata is an extremely large, simple, flattened, spatulate structure, about two- thirds the body length. All other species studied have a simple, unlobed penis. Refer to Harasewych and Kantor (1991:tables 2, 3) for a sunnnary of additional characters. Published information on marginelliform egg capsules was reviewed in Cxwvert (1986c and 1988e:42). Most marginelliform gastropods produce plano-convex or hemispherical egg capsules, but in Persicula cornea (La- marck, 1822) and Marginella goodalli (G. B. Sowerby I, 1825) they are stalked. In most cases, each egg capsule G. A. Coovert and H. K. Coovert, 1995 Page 65 contains a single embryo, but Pruniim apicimwi (Menke, 1828) and Dentimargo cairoma sometimes have a second embryo, while egg capsules of Volvarina avena contain about five embryos. Development is direct in all known cases. All marginelliform species have a paucispiral pro- toconch indicating direct development. Discussion: The division of marginelliform gastropods into two fundamentally different groups became obvious during the course of our research. These two groups were initially differentiated on gross radular features, but fur- ther research revealed differences in radular structure, anatomy, and internal columellar plications. In fact, these two groups can be differentiated on the basis of their internal shell whorls alone. These distinctly different groups are the cystiscids and the true marginellids. A brief summary of the differences follows. Cystiscids have a shell that possesses what we have termed cystiscid internal whorls (figures 2, 3), or modified cystiscid internal whorls in Graniilina (figure 4). These internal whorls are mostly resorbed by the animal, re- sulting in extremely thin, modified whorls. The anterior two or three columellar plications are reduced internally to a single, axially oriented, sharp edge. Within the ap- erture, the columella bears parietal lirae posterior to these plications, thus the columella is multiplicate In Gran- ulina, the posterior two columellar plications continue internally for a whorl or two and does not have a mul- tiplicate columella. Cystiscid radulae (figures 13-16) are long, narrow, and C-shaped, with over 80 and up to 200 or more rachidian plates (with lateral teeth in one genus). The narrow rachidian plates have few cusps and are weakly to distinctly arched. The two odontophoral car- tilages each fit into an anterolateral odontophoral carti- lage hood (modified in Granulina). which flank the rad- ula (figures 15, 16). The posterior end of the radula begins as a series of nascent rachidian plates. It is encased and secreted in a typical neogastropod radular sac, which extends beyond the end of the odontophore (figures 15, 35). True marginellids, on the other hand, have unmodified internal whorls (figure 1). The plications are continuous internally and the columella is not multiplicate. Margi- nellid radulae (figures 17-20) are a much shorter, usually broad, uniserial, J-shaped ribbon with 10 to 80 rachidian plates. These moderately to very broad, flat to weakly arched plates usually have many cusps. The subradular membrane is weakly expanded anterolaterally into mar- ginal cuticular flanges, which help hold the radula in correct alignment with the underlying, anteriorly fused odontophoral cartilages (figures 19, 20). The posterior end of the radula begins with only one or two preformed nascent rachidian plates which are secreted in the blind end of a buccal pouch. The odontophore extends past the posterior end of the radula (figure 19). After discerning the two fundamentally different groups of marginelliform gastropods, we searched for possible relationships among other neogastropod fami- lies. This revealed that all of the above cystiscid char- acters are shared with the Olividae. Thus, we feel that the cystiscids have a shared ancestry with that family, and not with the Marginellidae (see final Discussion sec- tion for more details). We have come to the conclusion that the cystiscids and the true marginellids are not close- ly related and must be considered as separate and distinct families. A systematic section, outlining the classification of these two groups, follows. SYSTEMATICS Key to the Families of Marginelliform Gastropods la. Columellar plications not continuous internally, at least anterior two columellar plications re- duced to one nearK' axiall)- oriented columellar edge within one full revolution internally; col- umella usually multiplicate with 3 (rarely 2) pli- cations, plus often up to 14 parietal lirae poste- riorly, but may have only 1 or 2 pseudocontin- uous plications posterior to the anterior 2 inter- nally reduced plications; internal whorls cystiscid or modified cystiscid type (figures 2, 3, 4) very thin and often partially missing due to partial resorption family CYSTISCIDAE lb. Columellar plications continuous internally, not reduced; columella with 2 to 6 plications, at least the first 4 continuing internally, the 5th and 6th often not continuing internally (and are thus parietal lirae); internal whorls unmodified type (figure 1) complete and of original thickness, without evidence of resorption family MARGINELLIDAE Family CYSTISCIDAE Stimpson, 1865:55 Diagnosis: Shell minute to large, white, uniformly col- ored, or patterned; surface smooth, sculptured, or axially costate; spire flat to immersed, or low to tall; protoconch paucispiral; lip thickened, smooth or denticulate; exter- nal varix present or absent; siphonal notch present or absent; posterior notch present or absent; columella mul- tiplicate, with combined total of 2 to 13 plications -I- parietal lirae or with 2 internally reduced columellar plications -I- 1 or 2 pseudo-continuous plications; internal whorls cystiscid or modified cystiscid type. Type 2, 3, or 4 animal; operculum absent. Cystiscid radula. Type 1, 2, 3, or 4. Mantle cavity with monopectinate ctenidium and bipectinate osphradium. Proboscis pleurembolic; jaws absent; typical radular sac present. Remarks: The family was erected by Stimpson (1865: 55) for a single included species based on unusual features of the head and radula. Subsequent authors included this group in the Marginellidae. Coan (1965:186) recognized this group as a subfamily, again based on unusual features of the head and radula, but did not include such genera as Gibberula (in part), Persicula. and Canaiispira. Rules of priority require this name to be used, which is un- fortunate because the type specimen of the type species Page 66 THE NAUTILI'S, Vol 109, Nos. 2 & 3 is lost (Coovert, 1986d) and the species is otherwise poorly known. We currently recognize 283 species in the Cys- tiscidae, of which 26 are undescribed. Key to the Recent Genera of the Family Cystiscidae la. Distinct siphonal notch present; spire low to im- mersed; shell minute to small 2 lb. Siphonal notch absent, anterior end rounded; spire immersed, low, or distinctly produced; shell mi- nute to large 3 2a. Adult shell completely lacking an external vari.x; spire low to nearly immersed, but never com- pletely immersed; shell usually white, uniformly colored in some, rarely with a few narrow spiral bands Gibberula 2b. Adult shell completely lacking an external varix or with an obscure to distinct, raised external varix; spire usually completely immersed, rarely very low; shell usually uniformly colored or with distinctive pattern of spots, tear-drop markings, longitudinal or oblique streaks or zig-zag lines; narrow spiral bands, if present, very numerous; rarely white, if so, with pattern Persicula 3a. External varix completely absent; columella mul- tiplicate, with combined total of 2 to 17 plications -I- parietal lirae; with cystiscid internal whorls. . .4 3b. External varix present, distinct, raised, usually strong; columella with 2 internally reduced an- terior plications + 1 or 2 pseudo-continuous pli- cations; with modified cystiscid internal whorls ... 8 4a. Posterior corner of aperture sharply, narrowly, and deeply channeled; aperture usually lirate, smooth in some; sutures often channeled or im- pressed, if filled with callus, then still distinct; spire distinctly produced Canalispira 4b. Posterior corner of aperture rounded, not chan- neled; aperture smooth or denticulate, not lirate; sutures callused over, indistinct; spire immersed or low 5 5a. .Shell with very strong, sharply crested axial cos- tae; strongly developed parietal callus "shield" present; spire sunken, but not immersed, with nuclear whorls visible; 5 columellar plications present; lip heavily thickened, flared posteriorly, not denticulate Extra 5b. Shell lacking strong axial costae, although may have very weak axial growth lines or very faint wrinkles; parietal callus wash often present, but lacking strongly developed parietal "shield"; spire produced or completely immersed, but not as above; combined total of 2 to 17 columellar pli- cations -t- parietal lirae; lip narrowly to strongly thickened, smooth or weakly denticulate 6 6a. Suture of last whorl expanding rapidly for final half whorl, then abruptly swept posteriorly just before lip, giving characteristic shape to shell; spire nearK' flat to low, not immersed; shell shape usually cylindrical, elliptic, or oblong Plesiocystiscus 6b. Suture of last whorl not unusually expanding or narrowing; spire low to completely immersed; shell shape rarely cylindrical, usually elliptic, ob- ovate to broadly obovate, or subtriangular 7 7a. Spire usually immersed; plications usually square in cross-section, often appearing excavated due to collabral parietal callus ridge, which intersects them just outside of aperture Crithe 7b. Spire usually at least slightly produced; plications usually rounded in cross-section, rarely appear- ing excavated, if so, not due to collabral parietal callus ridge Cystiscus 8a. Shell usually smooth, if sculptured, very obscure; spire immersed; shell elongate to broadly elliptic, obovate, subtriangular, or pyriform, not cylin- drical Granulina 8b. Shell with distinct, punctate sculpture; spire flat to immersed; shell broadly cylindrical and abruptly narrowed anteriorly to broadly subtrian- gular Pugnus Subfamily PLESIOCYSTISCINAE Coovert and Coo- vert, new subfamily Diagnosis: Shell minute to small, white, hyaline; last whorl rapidly expanded then lip abruptly swept poste- riorly giving characteristic shape; spire flat to low; lip thickened posteriorly, smooth, lacking lirae or denticu- lation; external varix absent; siphonal notch absent; pos- terior notch absent; columella multiplicate with com- bined total of 3 to 8 plications + parietal lirae; internal whorls cystiscid type. Type 2 animal; tentacles and si- phon moderately long; mantle translucent, in some spe- cies not readily extending over external shell surface; foot relatively narrow. Type 1 radula. Internal anatomy unknown. Genus Plesiocystiscus Coovert and Coovert, new genus (figures 3, 21, 43) Type species: Marginella jewettii Carpenter, 1857a; OD, herein (figure 43). Diagnosis: Same as for subfamily. Description: Shell (figures 3, 43) minute to small, (adult length 1.5-6.0 mm). Color white, hyaline; surface smooth, glossy. Shape cylindrical, elliptic, or obovate; weakly to distinctly shouldered. Suture rapidly descending on last half (if last whorl, then abruptly sweeping upward just before lip, giving characteristic shape to adult shell. Spire nearly flat to low. Aperture moderately narrow, broader anteriorly. Lip thin anteriorly, gradually thickening to- ward shoulder, smooth, lacking denticulation or lirae, external varix absent. Siphonal notch and posterior notch ab.sent. Shell often with a weak parietal callus wash or weak parietal callus deposits. (]olumella multiplicate, with combined total of 3-8 plications -l- parietal lirae, usually G. A. Coovert and H. K. Coovert, 1995 Page 67 occupying less than half to slightly more than half the aperture length. Shell with cystiscid internal whorls. External anatomy: 4 species studied. Type 2 animal, with moderately long tentacles; eyes at base of tentacles; siphon simple, moderately long; mantle translucent and barely visible, possibly not extending over external shell surface in some species; foot relatively narrow, about as wide as shell and l'/2 times shell length; animal variously marked with tiny dots of various colors, pattern of in- ternal mantle showing through translucent shell. Internal anatomy: Unknown. Radula (figure 21): 3 species studied. Type 1, triserial, with simple, thin, weak laterals that narrow to a single cusp. Ribbon long, narrow, composed of 66-103 rows of plates. Rachidian plates overlapping, very narrow (0.017- 0.028 mm wide), weakly arched, with 3-7 sharp cusps on posterior edge. Anterior edge of rachidian plate slight- ly concave, resulting in weakly U-shaped plates. Shell length; radular width ratio (based on rachidian plate width) = 159-200. Radular Index = 10.9-34.3. Included species: P. abbotti (Jong & Coomans, 1988), P. atomus (E. A. Smith, 1890), P. bubistae (Fernandes, 1987), P. cinereus (Jousseaume, 1875), P. consanguineus (E. A. Smith, 1890), P. gutta (Gofas & Fernandes, 1988), P. jansscni (Jong & Coomans, 1988), P. jewettii (Car- penter, 1857), P. josephinae (Fernandes & Rolan, 1992), P. larva (Bavay, 1922), P. palantirulus (Roth & Coan, 1968), P. polituhis (Dall, 1919), plus 4 undescribed spe- cies. Distribution and Habitat: Indo-Pacific (1 species), E Pacific (4 species), western Atlantic (5 species), W. Af- rican (6 species). Intertidal to 80 m. Fossil Record: Eocene of France, Miocene and Pliocene of Florida, and Pleistocene of California, to Recent. Nomenclature, Synonymy: Photographs of the syntypes of the type species, Marginella jewettii. were published by Palmer (1958:pl. 24, figs. 19-21). A lectotype was designated and illustrated in Coan and Roth (1966:pl. 51, fig. 66). The genus name is derived from the Greek plesios. near, in the sense of plesiomorphic, of characters near the ancestor, combined with Cystiscns. This indi- cates the relationship to (and previous inclusion in) the genus Cystiscus, type genus of the family. The triserial radula, found in many other neogastropod families, but unknown in any other cystiscids, is the most primitive in this family. Gender masculine. Remarks: This is the only genus of marginelliform gas- tropods (previously referred to as "Cystiscns") with a triserial radula. This group is considered to be the most primitive in the family based on the radula and possession of a generalized Type 2 animal. Radular and shell fea- tures, including presence of odontophoral cartilage hoods, possession of cystiscid internal whorls, and lack of an external varix, place this group in the Cystiscidae. Gofas and Fernandes (1988:19) described the mantle of P. gutta (Gofas & Fernandes, 1988) as "translucent and hardly visible when it extends over the shell." Roth and Coan (1968:66) observed that "the mantle was never extruded over the top of the shell of C. jewettii" and Behrens (1984:241) likewise described the mantle as not covering the shell in the same species. Additional obser- vations on this point would be useful. Subfamily CYSTISCINAE Stimpson, 1865:55 Diagnosis: Shell minute to small, white; spire immersed to low; surface smooth or axially costate; lip thickened, smooth or denticulate; external varix absent; siphonal notch absent; posterior notch absent; columella multi- plicate, with combined total of usually 2 to 8 plications + parietal lirae; internal whorls cystiscid type. Type 3 animal; mantle smooth, at least partially extending over external shell surface. Type 2 radula. Internal anatomy unknown. t Genus Topaginella Laseron, 1957 Topaginclla Laseron, 1957:288 Type species: Marginella octoplicata Tenison-Woods, 1877 t; OD (M) Diagnosis: Shell minute, pyriform, strongly narrowed anteriorly; spire low; lip thickened, strongly denticulate; external varix absent (?); distinct axial costae present; siphonal notch absent; columella multiplicate, with com- bined total of usually 8 plications + parietal lirae. Fossil Record: Miocene of Australia. Nomenclature, Synonymy: The type species, T. octo- plicata, was figured in Cotton (1949;pl. 18). Remarks: The type species seems to be unique. The multiplicate columella and lack of a siphonal notch place it near Cystiscus, but the pyriform shape, exserted spire, denticulate lip, and fine axial costae ("striated growth lines" of Cotton, 1949:218) render it distinct. The absence of an external varix needs to be confirmed. Genus Cystiscus Stimpson, 1865 (figures 9, 10, 22, 44) Cystiscus Stimpson, 1865:55 Euliginella Laseron, 1957:282 [TS: Marginella angasi Crosse, 1870; OD] Type species: C. capensis Stimpson, 1865 (non Mar- ginella capensis Krauss, 1848), = Marginella cystiscus Redfield, 1870 (nom. nov.); M (figure 44). Diagnosis: Shell minute to small, white, hyaline; spire immersed to low; lip thickened, smooth or weakly den- ticulate; external varix absent; siphonal notch absent; posterior notch absent; lacking collabral parietal callus ridge; columella multiplicate, with combined total of Page 68 THE NAUTILUS, Vol. 109, Nos. 2 & 3 49 Figures 43-5 1 . Shells of type species of cystiscid genera, ventral views. 43. Plesiocystiscus jeicettii (Carpenter, 1857) GAC. M1823, Pirates C;ove San Luis Obispo, California, intertidal. Length 5 3 mm. 44. Cystiscus cystiscus (Redfield, 1870). Holotype (destroyed). False Bay. Cape of Good Hope, South Africa, 37 m, on gorgonians. After Stimpson (1865:pl 8, fig. 2). Length 3.6 mm. 45. Cnthe atomaria (louid. 1860. ANSP 307869, 1.6 km W.S.W, of Pulau Blankang Malti. Singapore, Malaya. Length 2.0 mm. 46. Extra extra Jous.seaume, 1894. P. W. Clover colln., Perim, South Yemen. Length 0.9 nun. 47. Pcrsicula persicula (Linne, 1758). GAC G. A. Coovert and H. K. Coovert, 1995 Page 69 usually 2 to 8 plications + parietal lirae, 1st plication usually strong and raised. Type 3 animal; mantle smooth, at least partially extending over external shell surface Description: Shell (figure 44) minute to small (adult length 1.0-6.0 mm). Color white, hyaline; surface smooth, glossy. Shape rarely cylindrical, usually elliptic, obovate, or subtriangular; weakK shouldered. Spire completely immersed to low. Aperture narrow to broad, usually wid- er anteriorly. Lip slightly to distincth thickened, flared posteriorly in some species, smooth on inside edge to weakly denticulate, lacking lirae, external varix absent. Shell lacking a siphonal notch and posterior notch. Shell with weak parietal callus wash or weak parietal callus deposits in some species, but lacking collabral parietal callus ridge. Columella multiplicate, with combined total of 2-8 plications + parietal lirae, rarely up to 17 in which the posteriormost are denticles; one species with onl\ 1 plication. Plications usually occupying less than half the length of the aperture, but most of the aperture in some. Plications excavated just inside aperture in a few species, usually evenK rounded, first plication usualK' raised and very strong. Shell with CNStiscid internal whorls. External anatomy (figures, 9, 10): 13 species studied. Type 3 animal with eyes at side of head, usually on lateral swelling; mantle smooth, at least partially extending onto external shell surface, in some species nearly covering shell; foot relatively narrow, about as wide as shell and 1-1.5 X shell length; head and mantle usually uniformly colored, often bright red, orange, or yellow, or brown or black, internal mantle color pattern often showing through translucent shell. Internal anatomy: Unknown. Radula (figure 22): 14 species studied. Type 2, uniserial, ribbon long, narrow, composed of 80-200 plates. Ra- chidian plates overlapping (rarely separate), very narrow (0.009-0.036 mm wide), weakly to strongly arched, with 5-15 very strong cusps on posterior edge. Anterior edge of rachidian plate slightly concave, resulting in crescent- shaped plates. Shell length: radular width ratio = 94- 217. Radular Index = 10.7-25.7. Distribution and Habitat: Neozelanic (1 species). So Australian (21 species), Indo- Pacific (11 species), W. At- lantic (2 species). South African (23 species). Intertidal to 370 m. Fossil Record: Eocene of France, Miocene and Pliocene of Australia, to Recent. Nomenclature. Synonymy: The type species C. cijstis- cits is discussed and figured in Coovert (1986d) and here- in (figure 44). The radula of this species, along with three others, was reviewed in Coovert (1989b:5-6). SEMs of an additional eight Australian species (Dean Hewish, pers. comm), including C. angasi, the type species of Eitliginella. have been examined by us. All these species are considered congeneric based on similar radulae and shell characters. Euliginclla was considered synonymous by Coan (1965:190). Remarks: Originally described in its own family by Stimpson, apparently based on the radula and unusual features of the head, this genus again becomes the nom- inate genus of a now more clearly defined family. Genus Crithe Gould, 1860 (figure 45) [TS: M. nipponica Habe. 1951; Crithe Gould, 1860:384 Microviilina Habe, 1951:105 OD] Type species: C. atomaria Gould, 1860; M (figure 45) Diagnosis: Shell minute to small, white, hyaline; spire usually immersed, rarely low; lip thickened, smooth, lacking denticulation; external varix absent; siphonal notch absent; posterior notch absent; columella multi- plicate, with combined total of 6-8 plications -I- parietal lirae, plications usually excavated inside aperture due to collabral parietal callus ridge. Presumably a Type 3 an- imal. Description: Shell (figure 45) minute to small (adult length 1.6-3.2 mm). Color white, hyaline; surface smooth, glossy. Shape narrowly to broadly obovate or subtrian- gular; weakly to moderately strongly shouldered. Spire usually completely immersed, rarely low. Aperture nar- row, wider anteriorly. Lip moderately thickened, es- pecially posteriorly, thinner anteriorly, smooth on inside edge, lacking denticulation and lirae, external varix ab- sent. Shell lacking a siphonal notch and posterior notch. Shell usually with a distinct collabral parietal callus ridge that intersects plications. Columella multiplicate, with combined total of 6-8 plications -t- parietal lirae, usually occupying about two-thirds the aperture length. Plica- tions usually appearing excavated just inside aperture due to callus deposits from anterior end of collabral pa- rietal callus ridge. Shell with cystiscid internal whorls. External anatomy: Presumably a Type 3 animal; man- tle extension undetermined; generally uniformly colored, bright yellow, rose, or orange-red, or black with 2 zones of yellow on internal mantle. M1558, Dakar, Senegal, West Africa, in sand, low tide. Length 20.0 mm. 48. Gibherula onjza (Lamarck, 1822). Neotype, MHNG, Senegal. After Gofas (1990:f)g. 2). Length 6,5 mm. 49. Canalispira olivellaeformis Jousseaume, 1875. Lectotype, MHNP, locality unknown. After photos in Kilburn (1990:fig. 5). Length 4,2 mm, 50. Granulina isseli (G. & H, Neviii, 1875), GAG Ace. # 53-87, Red Sea coast of Egypt between El Ghurdaga and 80 km south of El Quseir. Length 1.6 mm 51. Ptignus parvus Hedley, 1896. GAG M2135, east of North Ft., Flinders Island, Bass Strait, Australia, 39°43'S, 148°36'E. Length 2.0 mm. Page 70 THE NAUTILUS, Vol. 109, Nos. 2 & 3 Internal anatomy: Unknown. Radula: 1 species studied (3 specimens). Type 2, uni- serial, ribbon long, narrow, composed of 189-220 plates. Rachidian plates overlapping, very narrow (0.007-0.008 mm wide), weakly arched, with 4 strong cusps on pos- terior edge. Anterior edge of rachidian plate slightly concave, resulting in an asv mmetrically V-shaped plate. Shell length: raduiar width ratio = 234-270. Radular Index = 47.2-55.0. Distribution and Habitat: Indo-Pacific (6 species). In- tertidal to 58 m. Fossil Record: Pliocene of Australia to Recent. Nomenclature, Synonymy: The type species of Crithe and Micwviilina, and their generic synonymy, were dis- cussed in Coovert (1986e and 1987c). The holotype of C. aiomaria was figured in Johnson (1964:pl. 8, fig. 12) and Coovert (1986e;fig. 1). Remarks: The columellar plications, which are square in cross-section and excavated due to the collabral pa- rietal callus ridge, and the usually immersed spire, serve to differentiate this group. The radulae are diagnostic. Genus Extra Jousseaume, 1894 (figure 46) Extra Jousseaume, 1894:98, 101 Type species: £. extra Jousseaume, 1894; M (figure 46) Diagnosis: Shell minute, white, semitranslucent; prom- inent axial costae present; spire sunken but not immersed; lip strongly thickened, smooth, lacking denticulation, flared posteriorly; siphonal notch absent; posterior notch absent; distinct parietal callus "shield" present; columella multiplicate, with combined total of 5 plications + pa- rietal lirae, plications slightly excavated inside aperture due to parietal callus deposits. Description: Shell (figure 46) minute (adult length 1.3- 1.4 mm). Color white, body whorl semitranslucent, sur- face with 17-20 prominent, sharply crested axial costae Shape obovate, truncate posteriorly; strongly shouldered Spire sunken but not immersed. Aperture narrow, slight- ly wider anteriorly. Lip very strongly thickened, es- pecially posteriorly, arched above apex and flared pos- teriorly, smooth on inside edge, lacking denticulation and lirae, apparently lacking an external varix. Shell lacking a siphonal notch and posterior notch. Shell with a distinct parietal callus "shield" that is free anteriorly with slight chink behind Columella multiplicate, with combined total of 5 ])licati(jns 4 parietal lirae occupying about % the aperture length. Plications slightly excavated just in- side aperture due to callus deposits from the parietal callus shield. Internal whorls unknown but presumed to be cystiscid type. External anatomy: Unknown. Internal anatomy: Unknown. Radula: Unknown. Distribution and Habitat: Indo- Pacific (1 species), re- corded only from the western Indian Ocean. Apparently shallow water. Fossil Record: K.nown only from the Recent. Nomenclature, Synonymy: See Coovert (1987d) for dis- cussion. Remarks: This monotypic genus is provisionally placed in the Cystiscinae based on the 5 excavated plications and lack of labial denticulation. The last strong axial costa has the appearance of an external varix, which, if present, might eliminate this genus from the Cystiscinae. The slightly sunken but non-immersed spire and the parietal "shield of this group appear to be unique. Subfamily PERSICULINAE Coovert and Coovert, new subfamily Diagnosis: Shell minute to large, white, uniformly col- ored, or patterned; spire immersed or low to tall; lip thickened, smooth or Urate; external varix present or absent; siphonal notch present or absent; weak to distinct posterior notch present; columella multiplicate with combined total of 3-13 plications -I- parietal lirae; in- ternal whorls cystiscid type. Type 4 animal; tentacles short to long, rarely absent; siphon short to long; eyes situated laterall> on head slightly below the base of the tentacles; mantle usually not extending over external shell surface. Type 3 radula. Odontophoral cartilages separate; valve of Leiblein present, with or without bypass tube; esophageal caecum absent; gland of Leiblein short, small, emptying directly into posterior end of esophagus; paired salivary glands ascinous or tubular, attached to esophagus just anterior to the valve of Leiblein, ducts attached to walls of esophagus; single accessory salivary gland pres- ent, ascinous or tubular; anal gland absent. Genus Persicula Schumacher, 1817 (figures 2, 11-16, 23, 35-37, 47) I'crsicula Scliunuiclier, 1817:235 Piwhijhathrun Claskoiii, 185;3:356[TS: P. rnarginclloidcuni Gus- koiii, 185.3, = Erato cijpracuidcs il. B .\(lanis, 1845; SD (MlTryon, 1885:270] Pcrsicida, suhg Ralncca Gray, 1857:37 [TS: /'. inlerrupta (La- marck, 1822), = Marginella intcrrupta Lamarck, 1822, = Volulci iulcrruplolineata Megerle von Miihifeld, 1816; Ml Type species: P. variabilis Schumacher, 1817, = Voluta persicula Linne, 1758: M (figure 47) Diagnosis: Shell small to large, uniformly colored, or white to colored and patterned, nuxlerately thick to thick- shelled; spire usualK inunersed; lip thickened, vveakK to strongly lirate; external varix present or absent; distinct siphonal notch present; posterior notch present; colu- mella nniltiplicatc with combined total of 4-13 plications G. A. Coovert and H. K. Coovert, 1995 Page 71 + parietal lirae. Type 4 anitnai; teiitatles long; siplion usually long; mantle apparently not extending over ex- terna! shell surface. Description: Shell (figures 2, 47) small to large (adult length 4.5-34 0 mm). CJolor white, tan, pinkish- or yel- lowish-tan, to gray, with pattern of spiral lines, rows of dots or tear-drop shaped markings, or longitudinal wavy or zig-zag markings which often form triangular peaks, less often uniformly Desh-colored or nearly white due to pattern reduction; surface smooth, glossy, few species with weak axial costae. Shape narrowly to broadly ob- ovate, or narrowly to broadly elliptic; weakly to mod- erately strongly shouldered. Spire usually completely im- mersed, rarely slightly produced. Aperture narrow, slightly wider anteriorly Lip moderately thickened, es- pecially posteriorly, thinner anteriorly, very weakly to strongly lirate, completely lacking an external varix or with strong to weak external varix Shell with a distinct siphonal notch and a very weak to very distinct posterior notch. Moderately-thick to thick-shelled, some species with distinct parietal or apical callus deposits, a few with a very strong parietal callus "shield." Columella multi- plicate, with combined total of 4-13 plications + parietal lirae usually occupying more than half the aperture length. Shell with c\stiscid internal whorls. External anatomy (figures 11, 12): 15 species studied. Type 4 animal with moderately long to long tentacles; siphon simple, moderately short to usually very long; mantle apparently not extending over external shell sur- face; foot narrow to moderately broad, about as wide as shell to considerably broader and about IVz times shell length; animal variously marked with tiny dots, spots, blotches, flecks, or streaks of opaque white, black, or various colors. Internal anatomy (figures 35-37): 2 species studied. As in family diagnosis. Valve of Leiblein without bypass tube, paired salivary glands ascinous, single accessory salivary gland ascinous. Radula (figures 13-16, 23): 14 species studied. Type 3, uniserial, ribbon long, narrow, composed of 80-209 plates. Rachidian plates overlapping, narrow (0.011-0.093 mm wide), moderately to strongly arched, with 6-14 sharp cusps on posterior edge, the central cusp usually the strongest. Anterior edge of rachidian plate strongly con- cave, resulting in \J- or V-shaped plates. Shell length: radular width ratio = 146-572. Radular Index = 6.7- 21.8. Distribution and Habitat: S. Australian (3 species), Indo- Pacific (11 species), E. Pacific (7 species), W. Atlantic (20 species), W. African (9 species), South African (1 species). Fossil Record: Eocene of France and Alabama, upper Oligocene and Miocene of W. Atlantic, to Recent. In- tertidal to 370 m. Nomenclature, Synonymy: The type species of Rabicea, V. interrtiptolineata, is congeneric with P. persicula, based on similar shell, radular (Coovert, 1989b:22), and external anatomical features. {]oan and Roth (1966282) discussed this synonymy. Pachybathron has been treated as a valid genus by many recent authors, including Coo- mans (1972), who discussed the group, its early confused familial placement, the type species, and its synonymy. The cited example in Adams and Adams (1853:194) does not constitute a type fixation. Tryon (1885:270) thus pro- vided the earliest type fixation by subsequent designa- tion. The features of the external anatomy, radula, and the brown triangular markings of the shell indicate a close relationship to some of the other small western Atlantic Persicuta. The parietal callus shield is considered to be no more than a species-group character. Thus, Pachybathron is here treated as a synonym. Subgeneric recognition may eventually be justified, but this would necessitate formal recognition of other species groups, a step considered unwise at this time. Remarks: Erroneously reported to have a Type 2 animal (Coovert, 1987g, 19871) based on strictly dorsal views, closer examination of several species revealed the split head typical of a Type 4 animal (figure 11). The two halves of the head are capable of closing together at will, thus appearing unsplit. The long siphon usually distin- guishes this group anatomically from the closely related Gibberiila. The shell of Persicula is usually patterned, often has a distinct external varix, and the spire is usually immersed. These conchological features serve to separate the two groups and are presented in the key to Recent genera. See Coovert (1987b) for a discussion of the Per- sicula cornea group. Genus Gibberula Swainson, 1840 (figures 48, 79) Gibberula Swainson, 1840:323 Grantila Jousseaume, 1875:167, 246 [TS: C. bei^soni (Reeve, 1865), = Marginclla bcnsoni Reeve, 1865; SD (M) Coan, 1965: 190] Kogoinea Habe, 1 951:) 03 [TS: Marginclla nuvenipruvincialis (Yokoyama, 1928), = Erato ii(>vcniprovincialis\oko\dma., 1928; OD] t Dcntiginella Laseron, 1957:288 [TS: Marginclla rnctula Cot- ton, 1949 t; OD (M)] Epiginella Laseron, 1957:279 [TS: £. ablita Laseron, 1957; OD] t Lataginella Laseron, 1957:288 [TS: Marginclla hitsoni Chap- man, 1921 t; OD (M)] Phyloginella Laseron, 1957280 [TS: P. coinprcssa Laseron, 19.57; OD] t Vetaginclla Laseron, 1957:288 [TS: Marginclla donia Cotton, 1949 t;OD{M)] Dilnculum Barnard, 1962:14 [TS: D. inopinaliun Barnard, 1962, = Marginclla bcnsoni Reeve, 1865, OD] Type species: G. zonata Swainson, 1840, = Vulvaria oryza Lamarck, 1822; M (figure 48) Diagnosis: Shell minute to medium size, white or col- ored, some species with spiral bands; spire low to medium height, not immersed; lip thickened, weakly to strongly Page 72 THE NAUTILUS, Vol, 109, Nos, 2 & 3 lirate, smooth in some species; external varix absent; dis- tinct siphonal notch present; posterior notch present but often weak; columella multiplicate with combined total of 3-8 plications + parietal lirae. Type 4 animal; ten- tacles short; siphon short; mantle not usually extending over external shell surface. Description: Shell (figure 48) minute to medium in size (adult length 1.8-8.5 mm). Color usually white, hyaline, less commonly tan, yellow, or orange, some species with spiral lines or bands or sutural band, rarely with oblique markings; surface smooth, glossy, rarely with fine, weak axial costae. Shape narrowly subcylindrical to usually elliptic, obovate, or subpyriform; weakly to strongly shouldered, some species with weakly raised sutures. Spire low to medium height, not immersed. Aperture narrow to moderately broad, wider anteriorly. Lip moderately thickened, especially posteriorly, thinner anteriorly, usu- ally weakly to strongly lirate but absent in some species, completely lacking an external varix. Shell with distinct siphonal notch and very weak to distinct posterior notch. Shell with apical callus deposits in some species, rarely with elongate collabral parietal callus deposits. Colu- mella multiplicate, with combined total of 3-8 plications + parietal lirae occupying Va to nearly the full aperture length. Shell with cystiscid internal whorls. External anatomy: 26+ species studied. Type 4 animal, with short tentacles, which are rarely absent; siphon sim- ple, short, barely extending above the dorsal edge of the siphonal notch; mantle apparently not extending over external shell surface or with a small lobe on the left side in some species; foot narrow to moderately broad, about as wide to slightly wider than shell and usually about IVi times shell length; some species with anterolateral edges of foot weakly to strongly raised to form "parapodia ; animal variously marked with tiny dots, spots, or blotch- es, but not striped or lineated, markings of opaque white, brown, yellow, orange, or green. Internal anatomy (figure 79): 1 species studied. As in family diagnosis. Valve of Leiblein with bypass tube; paired salivary glands tubular; single accessory salivary gland tubular. Radula: 21 species studied. Type 3, uniserial, ribbon long, narrow, of 80-153 plates. Rachidian plates over- lapping, narrow (0.008-0.042 mm wide), moderately to strongly arched, with 5-11 sharp cusps on posterior edge. The central cusp is usually the strongest. Anterior edge of rachidian plate strongly concave, resulting in U- or V-shaped plates. Shell length: radular width ratio = 118- 309. Radular Index = 9.1-21.9. Distribution and Habitat: Neozelanic (2 species), S. Aus- tralian (4 species), Indo-Pacific (26 species), E. Pacific (6 species), W. Atlantic (9 species), Mediterranean (10 spe- cies), W. African (25 species). South African (7 species). Intertidal to 3,300 m. Fossil Record: Paleocene and Eocene of France and W. Atlantic, Miocene and Pliocene of Italy, Australia, W. Atlantic, and Japan, and Pleistocene of W. Atlantic and California, to Recent. INomenclature, Synonymy: Gibberula onjza. type spe- cies of this genus, is valid and distinct from G. miliaria (Linne, 1758). Coan (1965:189) (questionably) and Dodge (1955:79-82) considered these two species synonymous. But Gofas (1990:117) clearly differentiated the two spe- cies based on shell and external anatomical characters, and designated a neotype (figure 48) and lectotype re- spectively. The geims Granula and its type species M. bensoni differ from Gibberula (type species G. oryza) only in size and very minor, species-level shell characters. Granula was synonymized under Gibberula by Coss- mann (1899:96) and Wenz (1943:141). Kogomea, like- wise differing only in minor characters, was synonymized by Roth and Coan (1971:579) and Rehder (1980:85). Diluctilum was discussed and synonymized by Kilburn and Rippey (1982:113, 214). Barnard (1962) described Dibicubin} and its type species simply because the radula was known, whereas other species "are merely names given to the shells of molluscs whose anatomy is unknown (op. cit., p. 14)." This is clearly untenable. The wishbone- shaped radula was an error in observation, likewise made by Ponder (1970:70, fig. 3F). This error was discussed in Coovert (1989b:12). Gofas (1989a) correctly figured a number of radulae from Gibberula, drawn from SEMs. The five Laseron (1957) genera were all based on species- level characters, and are here synonimized with Gibber- ula. The t\pe species of all these genera show a distinct siphonal notch. The type species of Phyloginella was figured by Kaicher (1992: #6211). Powell (1979:222) synonymized Lataginella under Kagomea (sic). Remarks: Gibberula cucullata Gofas and Fernandes, 1988, with strongly developed "parapodia," has a highly modified head. The bifurcated tips of the head taper into divergent anterior lobes. These tips can be interpreted as being either the anterior lobes of the head and thus lacking tentacles, or the tentacles themselves and lacking the anterior lobes. Based on the sharply pointed tips and relative position of the eyes, we interpret these tips as being the anterior lobes of the head, the animal lacking tentacles. Gofas and Fernandes (1988:25) suggested that this species "may deserve separate generic status." Gib- berula suhbulbosa (Tate, 1878) (based on unpubl. data ex. R. Burn and D. Hewish) has a similarly constructed head but apparently lacks the strongly developed "par- apodia." Until the external anatomy of the t>pe species of all related genus-group names are known, it is con- sidered unwise to place these species in a different genus- level group. This large, widely distributed group is well charac- terized by radular, external anatomical, and concholog- ical characters. See Persicula for additional discussion. Genus Canalispira Jousseaume, 1875 (figure 49) (.'(luaiisiiiru JoiisseaiiiiR', 187.5:168, 270 G. A. Coovert and H. K. Coovert, 1995 Page 73 Baroginella Laseron, 1957:286-287 [TS: B. infirma Laseron, 1957; OD] Type species: C. olivellaeformis Jousseaume, 1875; M (figure 49) Diagnosis: Shell minute to medium size, white, hyaline, rarely with faint pattern, cylindrical-biconic to obovate or obconic; sutures usually impressed to channeled; spire medium to tali; lip thickened, smooth or iirate; external varix absent; siphonal notch absent; deeply channeled posterior notch present; columella multiplicate with combined total of 3-6 plications + parietal lirae. Mod- ified Type 4 animal; tentacles absent; siphon not visible; foot split anteromedially, covering external shell surface anterolaterally. Description: Shell (figure 49) minute to medium in size (adult length 2.2-10.3 mm). Color usually white, hyaline, rarely with faint brown maculations, fine oblique lines, or weak axial streaks; surface smooth, glossy. Shape cy- lindrical-biconic to narrowly obovate or obconic; usually very weakly to weakly shouldered. Sutures distinctly to weakly channeled, deeply impressed, or completely cal- lused over. Spire medium to tall. Aperture narrow to moderately broad, wider anteriorly. Lip moderately thickened, especially posteriorly, thinner anteriorly, out- er edge incurved on posterior half, usually weakly to strongly Urate but smooth and lacking lirae in some spe- cies, completely lacking an external varix. Shell lacking a siphonal notch but with a sharply, narrowly, and deeply channeled posterior notch. Parietal callus wash usually confined to aperture. Columella multiplicate, with com- bined total of 3-6 plications -f- parietal lirae occupying half or more of the aperture length. Shell with cystiscid internal whorls. External anatomy: 2 species studied. Modified Type 4 animal; tentacles absent; triangular lobes of head pointed anteriorly; the small black eyes medially placed on each triangular lobe; siphon not visible; mantle extension un- determined; foot narrow, about as wide and long as the shell, split anteromedially, covering the anterior end of the shell laterally like "parapodia ; animal white or col- orless. (Largely based on unpubl. notes and drawings of an undescribed W. Australian species, ex. P. W. Clover). Internal anatomy: Unknown. Radula: 2 species studied. Type 3, uniserial, ribbon long, narrow, composed of 121-136 plates. Rachidian plates overlapping, narrow (0.016-0.033 mm wide), moderate- ly to strongly arched, with 7 sharp cusps on posterior edge. The central cusp is usually the strongest. Anterior edge of rachidian plate strongly concave, resulting in U- or V-shaped plates. Shell length: radular width ratio = 176-270. Radular Index = 17.3-19.4. Distribution and Habitat: So. Australian (1 species), Indo-Pacific (7 species), W. Atlantic (3 species). South African (2 species). Intertidal to 540 m. Fossil Record: Pliocene of Australia to Recent. Nomenclature, Synonymy: The lectotype of C. olivel- laeformis was figured by Kilburn (1990:fig. 5) and is illustrated here (figure 49). We studied the holotype of B. infirma and it is clearly congeneric, differing only in the degree of channeling of the sutures. We regard this character to be of species-level and synonymize Baro- gijiella with Canalispira. Remarks: The modified Type 4 animal and lack of a siphonal notch render this group distinct, but the Type 3 radula clearly places this genus in the Persiculini. More study is needed of living animals. Subfamily GRANULININAE Coovert and Coovert, new subfamily Diagnosis: Shell minute to small, white, hyaline; surface smooth or sculptured; spire flat to immersed; lip thick- ened, smooth or denticulate; external varix present; si- phonal notch absent; columella with 2 internally reduced columellar plications, plus 1 or 2 pseudo-continuous pli- cations; internal whorls modified cystiscid type. Type 2 animal; tentacles long, slender; siphon moderately long; mantle pustulose, extending over external shell surface; foot long, narrow. Type 4 radula. Internal anatomy un- known. Genus Granulina Jousseaume, 1888 (figures 4, 24, 50) Granulina Jousseaume, 1888:191 ? f Cryptospira (Cyprafo/ina) Cerulli-Irelli, 1911:231 [TS: Val- uta clandestina Brocchi, 1814 f; M] Mcrovia Dall, 192186 [TS: Merovia pyriformis (Carpenter, 1865), = Volutclla pyriformis Carpenter, 1865, = ? Mar- ginella margaritula Carpenter, 1857b; M) Microginella Laseron, 1957:280 [TS: Marginella anxia Hedley, 1909; OD] Gibberulina audi., non Monterosato, 1884 Type species: Marginella pygmaea Issel, 1869 {non Marginella pygmaea G. B. Sowerby II, 1846), = Mar- ginella isseli G. & H. Nevill, 1875 (nom. nov.); M (figure 50) Diagnosis: Shell minute to small, white, hyaline; spire immersed; lip strongly thickened, usually denticulate; external varix present; siphonal notch absent; collabral parietal callus ridge often present; columella with 4 strong plications that are often excavated inside aperture. Type 2 animal; tentacles long, slender; siphon moderately long; foot long, narrow; mantle pustulose, extending over ex- ternal shell surface. Description: Shell (figures 4, 50) minute to small (adult length 0.8-3.2 mm). Color white, hyaline; surface smooth, glossy, rarely with fine, weak axial costae or minutely pitted. Shape narrowly to broadly elliptic, obovate, sub- triangular, or pyriform; very weakly to weakly shoul- dered. Spire usually immersed. Aperture narrow to mod- erately laroad, noticeably wider anteriorly and slightly wider posteriorly. Lip moderately to greatly thickened, Page 74 THE NAUTILUS, Vol. 109, Nos. 2 & 3 especially posteriorly, usually distinctly denticulate, rare- ly smooth, with distinct external varix. Shell lacking a siphonal notch; posterior notch usually absent, rarely weakly developed. Shell usually with a weak to distinct collabral parietal callus ridge that often intersects colu- mellar plications. (Columella with 4 strong plications, third and fourth rarely weak, rarely with weak parietal den- ticles posterior to plications. Plications often appearing excavated inside aperture due to callus deposits from anterior end of collabral parietal callus ridge. Plications occujjying halt or less the aperture length. Shell with modified cysti.scid internal whorls. External anatomy: 13 species studied. Type 2 animal, with long, slender tentacles; eyes at base of tentacles; siphon simple, moderately long; mantle pustulose, ex- tending symmetrically over most of external shell sur- face; foot narrower to slightly wider than shell and about twice the shell length; animal variously marked with tiny dots, spots, or areas of concentrated pigmentation, mark- ings usually black, white, or yellow, but may be orange or turquoise. Internal anatomy: Unknown. Radula (figure 24): 12 species studied. Type 4, uniserial, ribbon long, narrow, composed of 90-159 plates. Ra- chidian plates usually completely non-overlapping, very narrow (0.006-0.012 mm wide), moderately arched, with 9 to 12 raised, clustered denticles located in staggered positions on dorsal surface, with a single (or asymmet- rically paired) strong central cusp protruding posteriorly. Anterior edge of rachidian plate weakly concave, re- sulting in vaguely V-shaped plates. The rachidian plates are asymmetrical and alternate as mirror images. Shell length: radular width ratio = 164-351. Radular Index = 8.2-13.7. Distribution and Habitat: So. Australian (3 species), Indo- Pacific (13 species), E. Pacific (3 species), W. At- lantic (12 species), Mediterranean (8 species), W. African (3 species), South African (3 species). Intertidal to 1,285 m (single record at 1,700 m). Fossil Record: Miocene to Pliocene of W. Atlantic, Pliocene ol Italy, Pleistocene of Japan and California, to Recent. Nomenclature, Synonymy: The type species and ge- neric synonymy oi Granulina was discussed in Coovert (1987h). Although the "problem of Granulina clandes- tina" was treated by Gofas (1992:5-6), with a neotype designated and figured (op. cit., fig. 3), several discre- pencies remain. Brocchi (1814:642-643, pi. 15, fig. 11) described and figured a shell with 3 cohmiellar plications, a slightly exserted spire, and a pyriform shape. This was considered to be a juvenile 1)\ ( iofas ( 1 992:5), but juvenile Granulina have four plications, an immersed spire, and an ovate or globose shape. More study is needed to resolve these discrepencies. The lectotype of V, pyrijormis. type species of Merovia, was figured by Cloan and Roth (1966: pi. 51, fig. 77). Further research is needed to clearly establish the synonymy of this species with M. margar- itula. We have studied syntypes ol M . anxia, type species of Microginella, and regard them to be referable to Granulina. Remarks: Absence of a siphonal notch and presence of a distinct external varix distinguish this genus from all other cystiscid genera except Piignus. Shell shape and usual lack of sculpturing will further characterize Gran- ulina. The very distinct Type 4 radula, modified cystiscid internal whorls, and Type 2 animal clearly distinguish these two genera as a separate subfamily. For a discussion of G. hadria (Dall, 1889), see Coovert (1988a). Genus Pugnus Hedley, 1896 (figure 51) Pugnus Hedley, 1896:105-106 Mar^incUopsis Ravav, 1911:241 [TS: M serrei Bavay, 1911; M] Type species: P. parvus Hedley, 1896; OD (M) (figure 51) Diagnosis: Shell minute, white, hyaline; surface mi- nutely sculptured; spire flat to immersed; lip thickened, smooth or denticulate; external varix present; siphonal notch absent; columella with 3 or 4 plications. Type 2 animal; tentacles long, slender; siphon moderately long; foot long, narrow; mantle pustulose, extending over ex- ternal shell surface. Description: Shell (figure 51) minute (adult length 0.75- 2.0 mm). C>olor white, hyaline; surface sculpture of ex- tremely fine, crossed spiral and axial threads leaving rows of quadrate or hexagonal pits in between, or with spiral or axial rows of beads or short, dashed ridges which are often serpentine or interconnected. Shape broadly cylin- drical to broadly subtriangular; weakly to strongly nar- rowed anteriorly, strongly shouldered. Spire flat to im- mersed. Aperture moderately narrow to broad, wider anteriorly and slightly wider posteriorly. Lip strongly thickened, especially posteriorly, distinctly denticulate or smooth, with distinct external varix. Shell lacking a siphonal notch and posterior notch. Often with a distinct parietal callus wash extending out of aperture. Columella with 3-4 plications, third and fourth often weak. Pli- cations occupying less than hall the aperture length. Shell with modified cysti.scid internal whorls. External anatomy: I species studied. Type 2 animal, with long, slemler tentacles; eyes at base of tentacles; siphon simple, moderately long; mantle pustulose, ex- tending symmetrically over most ot external shell sur- face; fool narrower than shell and about twice the shell length; animal variously marked with tiny reddish dots and si)ots on a translucent background. [Based on pers. obs. of pliotographs ot /'. serrei (Bavay, 1911), ex. Colin Redfern]. Internal anatomy: Unknown. G. A. Coovert and H. K. Coovert, 1995 Page 75 Radula: 2 species studied. Type 4, uniserial, ribbon long, narrow, of >124 plates. Racliidian plates usually com- pletely non-overlapping, very narrow (0.006 mm wide), moderately arched, with 11-12 raised, clustered denti- cles located in staggered positions on dor.sal surface, with a single (or as> mmetricaliy paired) strong central cusp protruding posteriorly. Anterior edge of rachidian plate weakly concave, resulting in vaguely V-shaped plates. The rachidian plates are asymmetrical and alternate as mirror images. Shell length: radular width ratio = 161- 242. Radular Index = 10.3. Distribution and Habitat: S. Australian (1 species), Indo- Pacific (1 species), W. Atlantic (2 species). Intertidal to 60 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: Marginellopsis is here syn- onymized with Piignus on the basis of very similar rad- ulae (pers. obs. of SEMs, ex. Dean Hewish) and similar shell sculpturing between the type species of the two groups. An undescribed species from Brazil has char- acters of both groups, and P. maesae Roth, 1972 is in- termediate in shell shape. Special attention should be paid to the distinctive surface sculpturing uniting them all. These differences are provisionally considered to be of specific value only. See Roth (1972:107) for remarks on differing familial placement. Remarks: For general comments, see remarks under Granulina. Family MARGINELLIDAE Fleming, 1828:328 Diagnosis: Shell minute to very large, white, uniformly colored, or patterned; protoconch paucispiral; lip thick- ened, smooth or denticulate; external varix present or absent; siphonal notch present or absent; columella with 2-6 plications; internal whorls unmodified type. Type 1 or Type 2 animal; operculum absent. Marginellid radula. Type 5, 6, modified Type 6, Type 7, 8, or 9, or non- radulate. Mantle cavity with monopectinate ctenidium and bipectinate osphradium. Proboscis pleurembolic; jaws absent; marginellid buccal pouch present, absent in non- radulate species; odontophoral cartilages fused anteriorly or both anteriorly and posteriorly; valve of Leiblein pres- ent or absent, with or without bypass tube; esophageal caecum present or absent; gland of Leiblein large, sac- culate, and emptying directly into posterior end oi esoph- agus, or with long, convoluted duct and a terminal bulb, emptying either into posterior end of esophagus or pass- ing through nerve ring and emptying into anterior end of proboscis; paired salivary glands ascinous or tubular, either attached to esophagus just anterior to the valve of Leiblein, contained within proboscis, or free, ducts either embedded in or attached to walls of esophagus, or free; single accessory salivary gland present or absent, ascinous or tubular; anal gland present. Remarks: We currently recognize a total of 483 species in the Marginellidae, of which 51 remain to be described. The two subfamilies were recognized in Coan (1965) but were not clearlv based on anatomical characters. Key to the Recent Genera of the Family Marginellidae la. Adult shell very large, over 50 mm in length; siphonal notch present, broad, weak to distinct; spire medium height, with large, often some- what bulbous protoconch; lip narrowly thick- ened, smooth, without denticulation 2 lb. Adult shell minute to large, rarely very large, if over 50 mm, then spire immersed or with strongly thickened, denticulate lip; spire im- mersed to tall, protoconch not unusually large; lip narrowly to strongly thickened, with or with- out denticulation 3 2a. Columella with 4 very strong, nearly transverse plications; 1st plication well posterior of anterior border of columella; with very distinct parietal callus deposit posteriorly; shell often somewhat dull and crazed or blistered Afrivoluta 2b. Columella with 2 strong, very oblique plica- tions; 1st plication bordering anterior end of columella; without distinct parietal callus de- posits; shell very smooth, glossy . . . Marginellona 3a. Columella with 4 strong plications occupying more than half the aperture length; if with weak "false 5th" plication, then with distinct poste- rior notch; spire low, medium, or tall; external varix distinct; often with weak to distinct si- phonal notch present 4 3b. Columella with 2-6 plications, those with 4 or fewer plications occupying half or less the ap- erture length; spire immersed, or low, medium, or tall; external varix distinct, weak, or absent; siphonal notch usually absent, if present, with more than 4 plications 12 4a. Shell with distinct to weak siphonal notch 5 4b. Shell completely lacking a siphonal notch, the anterior end rounded, or notch very weak and obscure 8 5a. Shell usually white or uniformly colored yel- lowish-white to \ellowish-orange, rarely with several obscure pale yellow spiral bands; Aus- tralian or Neozelanic distribution 6 5b. Shell usually with distinct and colorful pattern of spots or dots, numerous spiral lines, mottled, or with axial streaks or lines, or various com- binations; eastern Atlantic or western Indian Ocean distribution 7 6a. Posterior notch present, sharply defined; lip rapidly thickening posteriorly, narrowed slight- ly then widening again at posterior Vr. lip smooth, not denticulate; body whorl generally pyriform, strongly narrowed at anterior Vi; usually with heavy ventral callusing; spire whorls convex or shouldered, giving uneven profile; posterior end Page 76 THE NAUTILUS, Vol. 109, Nos. 2 & 3 of lip joins body whorl well below suture above; 4th plication not distinctly remote . . Amtroginella 6b. Posterior notch weak to absent; lip thickest me- dially, uniformly thickened throughout; lip smooth or weakly denticulate; body whorl ob- ovate, usually straight to convex at anterior Vi; spire whorls evenly contoured, essentially straight in profile; posterior end of lip joins body whorl at or slightly below previous suture; 4th plication often remote (i.e. farther separated than other 3) Mesoginella 7a. Shell with distinct to obscure axial costae; ap- erture usually broadest medially; lip strongly denticulate Glabella 7b. Shell smooth, completeK lacking axial costae; aperture usually broadest anteriorly; lip smooth or denticulate Marginella (part) 8a. Lip flared at shoulder, weakly to strongly alate; shell usually strongly narrowed anteriorly 9 8b. Lip not notably flared or alate; shell not usually strongly narrowed anteriorly 10 9a. With a strongly marked posterior notch in cor- ner of aperture; columellar plications appear excavated inside aperture due to callus deposits; weak "false 5th" plication present; lip strongly thickened, with distinct denticulation Protoginella 9b. With at most a weak, broadly rounded posterior notch in corner of aperture; columellar plica- tions rounded, not appearing excavated inside aperture; lacking "false 5th ' plication; lip mod- erately thickened, usually lacking denticula- tion, if present, usually weak Alaginella 10a. Shell either with intricate pattern of spiral and axial elements, or shell length > 15 mm Marginella (part) 10b. Shell usually white or uniformly colored; pat- tern, if present, consisting of only 1 to 4 narrow spiral bands; shell minute to medium in size, 1.9-12.0 mm in length 11 11a. Labial denticulation always present, subequal throughout; spire low to medium height; shell broadly obconic in shape; some species with weak to distinct axial costae Eratoidea lib. Labial denticulation sometimes absent, if pres- ent, with either a single strong posterior labial denticle, or posterior denticle distinctly stronger than others; spire medium to tall; shell usually narrowly biconic, rarely broadly biconic in shape; rarely with axial costae Dentimargo 12a. Spire immersed or nearly so 13 12b. Spire low to medium 20 13a. External varix extremely weak to absent; col- umella with 3 plications; shape narrowly cylin- drical Volvarina (part) 13b. External varix strong, distinct, if weak to absent, then with 4 columellar plications; columella with 3-6 plications; shape various 14 14a. Columella with 3 plications that are crowded anteriorly; shape cylindrical to narrowK' obo- vate 15 14b. Columella with 4-6 plications, that often oc- cupy half or more the aperture length; shape broadly elliptic to obovate, rarely cylindrical to narrowly obovate 16 15a. Without labial denticulation; bod\' whorl usually somewhat abruptK narrowed anteriorly Balanetta 15b. With fine labial denticulation in adult; body whorl gradual!} narrowed anteriorly Hydroginella (part) 16a. Columella with 4, 5, or 6 plications; if with only 4 plications, then lip completely lacking den- ticulation, shell uniformly colored or patterned, and shell moderately large to large Cryptospira (part) 16b. Columella with 4 plications; lip usualK' distinct- ly denticulate; if not denticulate, then shell white, and small to medium in size 17 17a. Columellar plications usually thin, sharp, an- teriorly crowded; if not, then lacking labial den- ticulation; Indo-Pacific distribution 18 17b. Columellar plications thick, moderately heavy, not crowded anteriorly; western Atlantic dis- tribution 19 18a. Labial denticulation strong, distinct; shell usu- ally with distinct color, lip and apical spot darker, rarely grayish-white; columella with 4 thin, sharp, anteriorly crowded plications; western Indian Ocean distribution Closia ISb. Labial denticulation usually absent, rarely weakly developed; shell \\ hite, lacking colora- tion; columella with 4 thin to moderately heavy plications occupying less than half the aperture length, crowded anteriorly in some; Australian or Neozelanic distribution Ovaginella 19a. Shell white; unpatterned or with 2 obscure, bro- ken spiral bands Prunum (part) 19b. Shell distinctly colored; distinctly patterned with spots or numerous spiral bands Bullata 20a. Columella with 3 plications that are crowded anteriorly; weak to strong collabral parietal cal- lus ridge present posterior to plications, the pos- terior end often appearing as "false 4th' pli- cation; lip with fine tlenticulation. . Hydroginella 20b. Columella with 2-6 plications that are not crowded anteriorly; usually lacking collabral parietal callus ridge; if present, then lip not denticulate, or with 2nd collabral callus ridge inside aperture and 4 plications; lip with or without denticulation, if present, usually strong ' 21 21a. Columella with 5-6 continuous plications; shell usually .solid gray, or gray with spiral lines or a distinct pattern of spiral lines interrupted by longitudinal streaks, but not white and un- marked; spire low, depressed; shell broadly el- liptic, obovate, pyriform, or subtriangular; me- G. A. Coovert and H. K. Coovert, 1995 Page 77 diiim-sized to large, length 6.2-46.0 mm, Indo- Pacific distribution Crtjptospira (part) 21b. Columella with 2-4 plications; if with more than 4 plications, then either (1) with a "false 5th" plication and shell narrow and cylindrical, (2) color pure white, opaque, with a weak siphonal notch, or (3) western Atlantic distribution; oth- erwise not as above 22 22a. Siphonal notch weak but present; Indo-Pacific, Australian, or Neozelanic distribution 23 22b. Siphonal notch completely absent; if weak but present, then West .\frican distribution; widely distributed 24 23a. Columella with 4 plications, 4th plication often remote; shell translucent white to white, with- out heavy ventral callus deposits; with distinct external varix; Australian or Neozelanic distribu- tion Mesoginella (part) 23b. Columella with 5-6 plications; shell opaque milky white w ith hea\\ ventral callus deposits; w ith or without external varix; Indo-Pacific dis- tribution Volvarina (part- nionilis Group) 24a. With "false 5th" plication present as a trans- verse denticle or weak parietal lira, neither ex- tending into aperture appreciably. rareK' ab- sent; lip denticulate when fully adult; shell cy- lindrical or subcylindical; distinct external varix present; shell colored and multi-banded or white; Indo-Pacific, Australian, or Neozelanic distribu- tion Serrate 24b. With 4 columellar plications; if 5th present, then continuous, not a "false 5th" or weak parietal lira; lip smooth or denticulate; shell \ariously shaped; if cylindrical or subcylindrical, then lip smooth, not denticulate; distinct external varix present or absent; widespread distribution .... 25 25a. Collabral parietal callus ridge present, although often weak, plus a collabral callus ridge present inside aperture, both just posterior to 4 strong columellar plications; lip with strong denticu- lation; external varix present, lip moderately to strongly thickened, especially at anterior third; shape obovate to subp\ riform; color translucent white or light orange; East African distribution Serrataginella 25b. Lacking collabral callus ridge inside aperture; if collabral parietal callus ridge present, then with onK 3 columellar plications and lip smooth; otherwise lacking above combination of char- acters 26 26a. Columellar plications 2, 3, or 4, thin and sharp; shell very thin, translucent or hyaline, color white to amber, usually unpatterned, rarely with faint yellowish-white spiral bands; external var- ix absent, if present, very weak; lip smooth, completely lacking denticulation Hyalina 26b. Columellar plications 3, 4, or 5, not unusually thin and sharp; if only 3, then joined by short collabral callus ridge posteriorly; shell color var- ious, shell often thicker, not unusually thin and translucent; external varix present or absent, if absent, columellar plications thicker; labial den- ticulation present or absent 27 27a. Shell with ventral and apical callus wash mi- nutely granulated; with distinct external varix; lip smooth, lacking denticulation; aperture broad; freshwater, SE Asian distribution Rivomarginella 27b. Shell lacking minute granulations ventrally and apically; without above combination of char- acters; not freshwater, widely distributed 28 28a. Labial denticulation present in man\ species, but may be absent; color uniformly white, cream, or gra\ , or spotted to streaked in some, but generally not with narrow darker bands alone (bands, if present, of pale background color interrupting darker shell color); columella with 4-5 plications; heavy ventral callusing common, especially callus pad near posterior commissure of lip; if shell white and lacking labial denticulation, then unpatterned, with 4 columellar plications, and western Atlantic dis- tribution Priinum 28b. Labial denticulation absent; color translucent white to orange or brown, usually with yellow, orange, brown, or reddish-brown narrow bands, which are darker than shell color, rarely axially streaked, not patterned with spots; columella with 3-4 plications; ventral callus deposits weak or absent Volvarina (part) Subfamily MARGINELLONINAE Coan, 1965:186, 191 Diagnosis: Shell very large; spire medium height; pro- toconch large; lip narrowly thickened, smooth, lacking denticulation; external varix present; siphonal notch pres- ent; columella with 2 or 4 strong plications. Type 1 an- imal; tentacles relatively short; eyes reduced or absent; siphon moderately long. Type 7 radula. Marginellid buc- cal pouch present; valve of Leiblein present, without bypass tube; esophageal caecum absent; gland of Leiblein large, sacculate, emptying directly into posterior end of esophagus; paired salivary glands tubular, free, ducts free along floor of proboscis; single accessory salivary gland absent. Genus Afrivoluta Tomlin, 1947 (figures 5, 6, 52) Afrivoluta Tomim, 1947:244 Type species: A. pringlei Tomlin, 1947; OD (M) (figure 52). Diagnosis: Shell very large, volute-shaped; spire me- dium height; protoconch large; lip narrowly thickened, smooth, lacking denticulation; external varix present; broad, distinct siphonal notch present; distinct parietal callus deposit present; columella with 4 strong, nearly Page 78 THE NAUTILUS, Vol. 109. Nos. 2 & 3 56 Figures 52-60. Sliclls of t\pc spocies ol marf^iiicllid genera, ventral \ lews, 52. Afrivohita prin^lci Tonilin, 19-17 i',\C. Ml 81 9, .'\j»ulhas Hank, Sonlli Alriea, trawled at 110 m Length 115.4 mm 53. Mar^ineUona gigas (Marten.s, 1904). Holotype of Sigalula pratasemis lU-hder, 19()7, USNM 237018, VV. of Pralas Reef, South China Sea, 20°37'N, 11,5°43'E. in 380 m, gray niud and sand bottom. After photos in Harasewych and Kantor (1991:fig. 2) and Weaver and DuPont (1970:pl. 40, fig. H, I). Length 54.1 mm. G. A. Coovert and H. K. Coovert, 1995 Page 79 transverse plications, 1st plication beginning well pos- terior of anterior end of columella. Type 1 animal; ten- tacles relatively short; eyes reduced; siphon moderately long; mantle smooth, extending at least partialK over external shell surface. Description: Shell (figure 52) very large (adult length 84.7-130.9 mm). Color tawny to cinnamon or pinkish- tan. Relatively thin-shelled, surface smooth and semi- glossy with evident growth lines, often crazed or with weak to distinct bubbles or blisters in shell surface. Shape oblong, volute-shaped, broadest at about posterior third, narrowed anteripe, Katow. New Guinea, mud bottom 13 m .After Hediey (1901:pi 16, fig 5). Length 6 4 mm. 58. Alaginella ochracea (Angas, 1871). GAG Ace. # 28-89, Little Bay, New South Wales. Length 3,5 mm. 59. Austroginella nmscaria (Lamarck, 1822). GAG M12.32, Victoria, Australia, on beach Length 15.3 mm. 60. Mesoginella lurbinata (G. B. Sowerby II, 1846). GAG MI655, Port Stephens, New South Wales. Length 7.2 mm. Page 80 THE NAUTILUS, Vol. 109, Nos. 2 & 3 long; mantle extension undetermined; foot broad, about 2 X shell width, slightly longer than shell (Harasewych and Kantor, 1991:10). Internal anatomy (figure 80): 1 species studied. Same as for subfamily. Radula (figure 28): 1 species studied. Type 7, uniserial, ribbon short, very broad, composed of 56-80 plates. Ra- chidian plates overlapping, very broad (1.6-2.5 mm wide), nearly flat, with numerous (58-85) sharp cusps along posterior edge. Anterior edge of rachidian plate generally straight, resulting in elongate, rectangular "comb-like" plates. Shell length: radular width ratio = 43-63. Radular Index = 0.9-1.0. Distribution and Habitat: Indo- Pacific (1 species). Oc- curs from the eastern Indian Ocean off the Nicobar Is- lands and the South China Sea. Recorded from depths of 380 to 1,280 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: The type species of Sigaluta is a synonym of M. gigas, the type species of Margi- nellona, thus synonymizing these two groups. See dis- cussion in Harasewych and Kantor (1991). The holotype of Sigaluta pratascnsis is figured herein (figure 53) and in color in Weaver and DuPont (1970:pl. 40, figs. H, I) and Abbott and Dance (1982:220). Remarks: Harasewych and Kantor (1991) reviewed this monotypic genus. This is the largest marginellid known. The smooth and very glossy shell, 2 very oblique colu- mellar plications, and lack of a parietal callus deposit serve to separate the type species from Afrivohita prin- glei, the only marginellid species with which it is likely to be confused. Subfamily MARGINELLINAE Fleming, 1828:328 Diagnosis: Shell minute to very large, white, uniformly colored, or patterned; lip thickened, smooth or dentic- ulate; external varix present or absent; siphonal notch present or absent; columella with 2-6 plications. Type 2 animal; tentacles long to very long, slender; eyes at base of tentacles on slight swelling; siphon moderately to very long; mantle smooth, pustulose, or papillose, usually at least partially extending over external shell surface. Type 5, 6, modified Type 6, Type 8, or 9 radula, or non- radulate. Marginellid buccal pouch present, absent in non-radulatc species; odontophoral cartilages fused an- teriorly or both anteriorly and posteriorly; valve of Lei- blein present or absent, if present, with bypass tube; esophageal caecum present or ab.sent; gland of Leiblein with long, convoluted duct and a terminal bulb, emp- tying either into posterior end of esophagus or passing through nerve ring and emptying into anterior end of proboscis; paired salivary glands a.scinous or tubular, ei- ther attached to esophagus jusl anterior to the valve of Leiblein, contained within proboscis, or free, ducts either embedded or attached to walls of esophagus, or free; single accessory salivary gland present or absent, ascinous or tubular. t Genus Myobanim Sohl, 1963 Mijoharum Solil, 1963:750-751 Type species: M . laevigatum Sohl, 1963 f; OD (M) Diagnosis: Shell moderately large, surface smooth, glossy spire medium to tall; sutures callused over; aperture broad lip thickened, smooth; external varix weakly developed weak siphonal notch present; strong, narrow posterior notch present; columella with 2 widely-spaced plications, the anterior one bordering the anterior end of the col- umella. Fossil Record: Late Cretaceous (Upper Maestrichtian) of Mississippi and Georgia. Nomenclature, Synonymy: Photographs of the holotype were figured in Sohl (1963:pl. 90, figs. 19, 20) and Sohl (1964:pl. 44, figs. 15, 16). Remarks: Apparently monotypic. The two widely- spaced columellar plications are unusual. Although orig- inally placed in the Volutidae, Ponder (1973:331) sug- gested that this genus might belong in the Marginellidae. Petuch and Sargent (1986:10) mentioned Myobaruin as a possible ancestor to the Olividae. The general shell shape, glossy, callused surface, anterior plication bor- dering the anterior end of the columella, and the blunt, paucispiral protoconch all indicate a marginellid. Based on these conchological features, this genus is here in- cluded in the Marginellidae, subfamily Marginellinae. Placement to tribe must await further, extensive study of the fossil fauna. Tribe AUSTROGINELLINI Coovert and Coovert, new tribe Diagnosis: Shell minute to moderately large, white, rarely lightly colored or obscurely banded; spire im- mersed to tall; lip thickened, smooth or denticulate; ex- ternal varix present; siphonal notch weak or absent; col- umella with 2-4 plications occupying more than half to less than half the aperture; some species with a "false 4th" or "false 5th plication. Type 2 animal; siphon mod- erately long to very long; mantle smooth, pustulose, or papillose, extending over external shell surface. Type 5, modified Type 6, or Type 8 or 9 radula. Marginellid buccal pouch present; odontophoral cartilages fused an- teriorly; valve of Leiblein present, with bypass tube; esophageal caecum absent; gland of Leiblein with a ter- minal bulb and a long, convoluted duct emptying into posterior end of esophagus; paired salivary glands tu- bular, attached to esophagus just anterior to the valve of i>eiblein, ducts embedded in walls of esophagus; single tubular accessory salivary gland present or absent. G. A. Coovert and H. K. Coovert, 1995 Page 81 "Serrata Group" Diagnosis: Shell small to medium, white to translucent, uniformly colored or banded in some species; spire im- mersed or low to medium; lip thickened, finely to coarse- ly denticulate, rarely smooth; external varix present; si- phonal notch absent; some species with collabral parietal callus ridge; columella with 3 or 4 plications occupying less than half the aperture length, often with "false 4th" or "false 5th plications. Type 2 animal; siphon mod- erately long; mantle smooth or pustulose, extending over external shell surface. Modified Type 6, or Type 8 or 9 radula. Anatomy as in tribe, single tubular accessory salivary gland present. t Genus Conuginella Laseron, 1957 Connginclla Laserun, 1957;2S8 Type species: Marginella inermis Tate, 1878 f; OD (M) Diagnosis: Shell medium size, conical, strongly but evenly narrowed anteriorly; spire low; lip narrowly thick- ened, denticulate, produced or angulate posteriorly; ex- ternal varix present; siphonal notch absent; columella with 4 plications occupying less than half the aperture length. Fossil Record: Miocene of Australia. Nomenclature, Synonymy: This group was placed by Coan (1965:190) as a subgenus of Serrata. Remarks: Related to Exiginella, a fossil group synon- ymized below with Serrata. Conuginella differs in hav- ing only 4 columellar plications, a distinctive conical shape, and a produced posterior corner of the lip. Shared features are the presence of an external varix, denticulate lip, and slender shell shape. Genus Serrata Jousseaume, 1875 (figures 26, 54) Serrata JoiLSseauine, 1875:167, 230 Haloginclla Laseron, 1957:284 [TS: Marginella nwstelina (Aii- gas, 1871), = Ilt/alina (\'i)lvarin(i) mtistelina Angas, 1871; OD] t Exiginella Laseron, 1957:289 [TS: Marginella winteri Tate, 1878 t: OD] Type species: Serrata .serrata (Gaskoin, 1849), = Mar- ginella serrata Gaskoin, 1849; T (figure 54) Diagnosis: Shell small to medium, white to brown, often banded, usually cylindrical; spire low to medium; lip thickened, finely to coarsely denticulate, rarely smooth; external varix present; lacking a siphonal notch; lacking parietal callus deposits and ridge; columella with 4 strong plications plus a weak "false 5th" occupying less than half the aperture length. Type 2 animal; siphon mod- erately long; mantle smooth or pustulose, extending over external shell surface. Modified Type 6 radula. Description: Shell (figure 54) small to medium in size (adult length 3.6-13.0 mm). Color white to grayish-white or brown, often with prominent brown spiral bands, or rarely pale orangish-yellow bands; surface smooth, glossy. Shape cylindrical to narrowly elliptic, narrowly subpy- riform in one species; weakly to strongly shouldered. Spire low to medium height, rarely very low and obscure. Aperture narrow, usually wider anteriorly. Lip weakly to strongly thickened, finely to coarsely denticulate, rare- ly smooth, with a weak to distinct external varix. Shell without a siphonal notch or a posterior notch. Shell with a thin parietal callus wash, lacking large callus deposits and collabral parietal callus ridge. Columella with 4 strong, continuous plications, plus a weak to strong pa- rietal lira or "false 5th" plication, which combined oc- cupy slightly less than half, but more than one-third, the aperture length. Internal whorls presumed unmodified. External anatomy: 2 species studied. Type 2 animal; tentacles long, slender; siphon moderately long; mantle smooth or pustulose, covering shell symmetrically; foot slightly wider than shell, about 1 V2 X shell length; animal colored with cream, orange, or buff. See summary in Coovert (1987g:13, 20) as Haloginella. Internal anatomy: 1 species studied. Same as for "Ser- rata Group. Radula (figure 26): 8 species studied. Modified Type 6, uniserial, ribbon short, broad, of 13-35 plates. Rachidian plates overlapping, thin, fragile, broad (0.052-0.120 mm wide), nearly flat, with numerous (22-59) sharp cusps along sinuous posterior edge. Anterior edge of rachidian plate generally straight, resulting in elongate, rectangular "comb-like" plates. Shell length: radular width ratio = 59-117. Radular Index = 0.4-1.6. Distribution and Habitat: Neozelanic (2 species), S. Aus- tralian (7 species), Indo-Pacific (3 species). Intertidal to 370 m. Fossil Record: Middle Oligocene of New Zealand {fide Powell, 1979:217), Miocene of Australia, to Recent. Nomenclature, Synonymy: The type of M. serrata is figured by Kaicher (1981:#2690) and herein (figure 54). Synonymy of Serrata and Haloginella is based on radular and conchological similarities. The radula of Australian S. mustelina, type species of Haloginella, has been stud- ied (pers. obs. of SEMs, ex. Dean Hewish) and compared with radulae from S. translata (Redfield, 1870), an un- doubted congener of S. serrata, type species of Serrata. These radulae and shells compare very favorably, and the generic groups they represent are herein considered synonyms. For discussion of Haloginella, see Coovert (1987f). Exiginella is synonymized based on concholog- ical similarity to the other two groups, including a fairly strong 5th columellar plication, denticulate lip, and an external varix. Remarks: Haloginella and Serrata had been considered distinct until we extracted radulae of S. translata. The Page 82 THE NAUTILUS, Vol. 109, Nos. 2 & 3 shared possession of a modified Type 6 radula, shell with a weak "false 5th" plication, lack of a collabral parietal callus ridge, and the usually denticulate lip are diag- nostic. (]oan (1965:190) considered Haloginella a sub- genus of Volvarina and was subsequently followed by others. Anatomical features described by Fonder (1970) clearly place this group, along with Mesoginella and Austroginella, in a separate tribe, based especially on the presence of a valve of Leiblein, the absence of an esophageal caecum, and the emptying of the duct from the gland of Leiblein into the esophagus posterior to the nerve ring. Genus Serrataginella Coo vert and Coovert, new genus (figures 29, 55) Type species: "Marginella" apryi Clover, 1974; OD (M), herein (figure 55) Diagnosis: Shell medium-sized, translucent white to light orange, unhanded, broadly obovate to subpyriforrn; spire low; lip strongly thickened, coarsely denticulate; external varix present; lacking a siphonal notch; with collabral parietal callus ridge plus parallel callus ridge inside ap- erture; cohnnella with 4 strong plications occupying about one-third the aperture length, lacking "false 5th ' pli- cation. Type 8 radula. Description: Shell (figure 55) medium in size (adult length 8.0-9.9 mm). Color translucent white to light or- ange; surface smooth, glossy. Shape broadly obovate to subpyriform, moderately strongly shouldered. Spire low. Aperture narrow, usually wider anteriorly, narrowest medially due to incurving of lip. Lip moderately to strongly thickened, especially anterior third, coarsely denticulate in adults, denticulation absent in subadults, first developing anteriorly, with a strong external varix. Siphonal notch absent; posterior notch absent. Shell with a weak but distinct collabral parietal callus ridge just outside aperture, plus weak to distinct collabral callus ridge well within aperture, l)oth most distinct just pos- terior to plications. Columella with 4 strong, continuous plications, lacking parietal lira or "false 5th ' plication. Plications occupying about '/s aperture length. Internal whorls presumed unmodihed. External anatomy: 1 species studied. Animal reported to be light orange in color ((Mover, 1974:215). Internal anatomy: Unknown. Radula (figure 29): 1 species studied. Type 8, iniiserial, ribbon short, broad, of .38 [)lates. Kacliidian plates over- lapping, broad (0.191 mm wide), nearly llat, with 9-10 sharp cusps along posterior edge. The entire posterior edge, including all of the edges of the main cusps, with numerous (total of ca. 70) small, subordinate cusps, giving the main cusps a serrated ajipearance. Anterior edge of rachidian plate slightly indented medially, resulting in a very sliallowly V-shaped plate. Shell length: radular width ratio = 42. Radular Index = 4.0. Distribution and Habitat: Indo- Pacific (1 species). Re- corded from E. African coast. Intertidal to 30 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: A photograph of the holo- type of the type species was figured by Clover (1974:pl. 8, fig. 6) and Kaicher (1981:#2629). Name derived from Serrata, valid marginellid genus to which this new genus is believed to be allied, and L. serrata, toothed like a saw, serrated, in reference to the serrated cusps of the radula, combined with -ginella, adopted from Margi- nella. Gender feminine. Remarks: This monotypic genus is characterized by the very distinctive radula, possession of a collabral callus ridge on the parietal area as well as inside the aperture, plus the 4 strong columellar plications and strong labial denticulation. Placement is based on conchological sim- ilarities to Serrata and Hydroginella. See Coomans (1975) for further information. See Key for further differenti- ation. t Genus Stronihoginella Laseron, 1957 Stromhoginella Laseron, 1957:289 Type species: Marginella erassidens Chapman and C;respin, 1928 f; OD (M) Diagnosis: Shell medium size, broadly biconic, strongly narrowed and acute anteriorly; spire medium height; aperture very narrow; lip thickened, denticulate; exter- nal varix present; siphonal notch absent; columella with 4 plications crowded anteriorly. Fossil Record: Pleistocene of Australia. Nomenclature, Synonymy: Coan (1965:189) placed this group as a subgenus of Marginella. but the anteriorly crowded plications indicate little relationship. Remarks: The broadly biconic shell shape strongly nar- rowing anteriorly and the very narrow aperture are ap- parently unique features of this monotypic genus. The ilenticulate lip and general shape indicate placement in Austroginellini, "Serrata Group, whereas the 4 anteri- orly crowded plications indicate close relationshij) to Hy- droginella. Genus Hydroginella Laseron, 1957 (figures 30, 56) llililrofiinrlla La.scroii, 1957:284 Nci>li)M>ncll(i Laseron, 1957:28.'! |'I'S: M . fa.sriciild Laseron, 1957; ODl rillargiiiella (laljtiel, 1962:197 (TS: Marginella colurimaria Hedley & May, 1908; OD (M)] Type species: //. dispersa La.seron, 1957; OD (M) (fig- ure 56) Diagnosis: Shell small to medium, white to orangish- brown or amber, hyaline or translucent, rarely banded, G. A. Coovert and H. K. Coovert, 1995 Page 83 usually cylindrical, obovate, or subpyriforni; spire im- mersed or low to medium; lip thickened, usually finely denticulate, rarely smootli; external varix present; lack- ing a siphonal notch; collabral parietal callus ridge usu- ally present; columella with 3 plications plus a weak "false 4th' occupying less than 'A tiie aperture length. Type 2 animal; siphon moderateK long; mantle extension undetermined. Type 9 radula. Description: Shell (figure 56) small to medium (adult length 3.0-10.3 mm). Color white to orangish-brown to amber, hyaline or translucent, one species with 3 broad pale orange spiral bands; surface smooth, glossy. Shape narrowly to broadly cylindrical or obovate to subpyri- form, usually strongly narrowed anteriorly; weakly to strongly shouldered. Spire immersed or low to medium height. Aperture usually narrow throughout, some spe- cies moderately broad, oiten wider anteriorly or narrow- est medially due to incurved lip. Lip moderately to strongly thickened, thickest medially or at anterior third, usually finely denticulate in adults, denticulation often weak or absent medially, rarely completely absent, with weak to strong external varix. Siphonal notch absent; posterior notch absent. Shell with weak to strong collabral parietal callus ridge beginning just posterior to plications, appearing at that point as "false 4th plication, ridge rarely absent. Columella with 3 continuous plications crowded anteriorly, occupying 'A or less of aperture length. Internal whorls unmodiHed. External anatomy: 2 species studied. Type 2 animal; tentacles long, slender; siphon moderately long; mantle extension undetermined; foot about as wide as shell, slightly longer, some species with opaque white spots. See Bouchet (1989:79, fig. 2). Internal anatomy: Unknown. Radula (figure 30): 4 species studied. Type 9, uniserial, ribbon very short, narrow, greatly reduced, of 10-30 plates. Rachidiaii plates weak, overlapping, narrow (0.019-0.032 mm wide), nearly flat, with few (4-7) cusps along posterior edge. Anterior edge of rachidian plate straight, resulting in subquadrate plates. Shell length: radular width ratio = 230-391. Radular Index = 2.3- 7.5. Distribution and Habitat: S. Australian (4 species), Indo- Pacific (13 species). Intertidal to 550 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: Bouchet (1989) discussed Hydrogi7iella and Neptoginella and figured the holo- types of both type species (note that his captions of figs. d and e were reversed). Although conchological similar- ities were discussed, these genera were not synonymized. These groups, plus Pillarginella, are here considered con- generic based on distinctive conchological features, es- pecially the presence of a collabral parietal callus ridge and only 3 columellar plications. The very distinctive radula is known from four species (Bouchet, 1989:fig. 3; Hewish, 1990:fig. 2A), including H. aihimnaria (pers. obs. of SEMs, ex. Dean Hewish), the type species of Pillarginella. The principle of first reviser (ICZN Art. 24) is employed in choosing Hydroginella over Nepto- ginella. These 3 groups form a very distinctive assem- blage of species. Remarks: Bouchet (1989) reported on the parasitism of sleeping fish by H. caledunica (Jousseaume, 1877). The very unusual, highly reduced radula, characteristic of this group, suggests a similar behavior in the other spe- cies. "Austroginella Group" Diagnosis: Shell minute to moderately large, white, rarely lightly colored or obscurely banded; spire im- mersed to tall; lip thickened, smooth or denticulate; ex- ternal varix present; siphonal notch weak or absent; col- umella with 2-4 plications occupying more than half to less than half the aperture, some species with a "false 5th ' plication. Type 2 animal; siphon long to very long; mantle smooth, weakly pustulose, or papillose, extending over external shell surface. Type 5 radula. Anatomy as in tribe, single accessory salivary gland absent. t Genus Mioginella Laseron, 1957 Mioginclla Laseron, 1957:287 Type species: Marginella regiila Cotton, 1949 f; OD (M) Diagnosis: Shell medium size, broadly biconic, nar- rowed anteriorly; spire tall; shoulders angulate or cari- nate; lip thickened, denticulate; external varix present; weak siphonal notch present; broad posterior notch pres- ent; columella with 4 plications, plus a weak 5th, occu- pying more than half the aperture. Fossil Record: Eocene of Australia. Nomenclature, Synonymy: This genus was placed as a subgenus of Marginella by Coan (1965:189). The fifth plication is not present in Marginella s. str. and the re- semblance is superficial. Remarks: We consider this apparently monotypic genus to be ancestral to Protoginella based on the presence of a weak 5th columellar plication, tall spire, and weak posterior notch. Laseron (1957:287) considered this genus to be ancestral to Carinaginella based on the carinate shoulders. Carinaginella is synonymized here with Al- aginella because we regard carinate shoulders to be con- vergent and not expressing a close relationship to mem- bers of Alaginella. which also has 4 columellar plications. Cotton (1949:218) stated that M. regula was related to M. muscaroides Tate, 1878, a species we consider to be clearly in Austroginella based on the distinctively shaped lip and presence of a strong siphonal notch, and thus not closely related to Mioginella. Page 84 THE NAUTILUS, Vol. 109, Nos. 2 & 3 Genus Protoginella Laseron, 1957 (figure 57) Protoginella Laseron, 1957:285 Type species: Marginella lavigata Brazier, 1877, = Marginella (Prunum) lavigata Brazier, 1877; OD (figure 57) Diagnosis: Shell small to medium, white, biconic or subpyriform, strongly narrowed anteriorly; spire medi- um height; aperture narrow; lip strongly thickened, strongly denticulate, alate posteriorly; external varix present; posterior notch present; siphonal notch absent; columella with 4 strong plications that are excavated inside aperture, plus a "false 5th"; plications occupying more than half the aperture. Description: Shell (figure 57) small to medium in size (adult length 4.4-7.0 mm). Color white; surface smooth, glossy. Shape biconic, subpyriform, strongly narrowed anteriorly, weakly to strongly shouldered. Spire medium height. Aperture narrow throughout. Lip strongly thick- ened, posterior corner strongly alate, strongly denticu- late, with distinct external varix. Shell without siphonal notch, but anterior end somewhat truncate; posterior notch present at posterior corner of lip. Extent of ventral callusing undetermined. Columella with 4 strong con- tinuous plications excavated inside aperture, plus weak "false 5th" plication or parietal lira; plications occupying 4/5 aperture length. Internal whorls presumed unmod- ified. External anatomy: LInknown. Internal anatomy: Unknown. Radula: 1 species studied. Type 5, uniserial, ribbon short, broad. Rachidian plates barely overlapping, mod- erately broad (0.028 mm wide), weakly arched, with 14- 15 strong cusps on posterior edge. Central cusp strongest. Anterior edge of rachidian plate moderately concave, resulting in rectangular to chevron-shaped plates. Shell length: radular width ratio = 160. Distribution and Habitat: Indo-Pacific (2 species). Re- corded from 13 to 97 m. Fossil Record: Eocene and Pliocene of Australia, to Recent. Nomenclature, Synonymy: A syntype of the type spe- cies is figured by Kaicher (1981:#2644) and herein (fig- ure 57). Hedley (1901:123) emended the name to M. laevigata, an unjustified emendation. No evidence in the original publication itself exists of "clear evidence of an inadvertent error," and incorrect transliteration is not to be considered an inadvertent error [ICZN Art. 32(c)(ii)]. Thus, there is no homonymy with Marginella laevigata Eichwald, 1830. The syntype of M. haudinensis Smith, 1899, a synonym, was figured by Kaicher (1981 :#2622), Some authors have synonymized M. valida Watson, 1886, which is an Alaginella and very distinct from P. lavigata. Remarks: Laseron's original concept included M. gem- inata Hedley, 1912. This was based on the presence of a denticulate lip, a character variable in Alaginella, in which this species is now placed. This restricted concept of Protoginella is based on a strong posterior notch, col- umellar plications appearing excavated due to callus de- posits, presence of a weak "false 5th" plication, and a strongly thickened, denticulate lip. t Genus Niidifaba Eames, 1952 Marginella. subg. Niidifaba Eames, 1952:122 Type species: Marginella (N.) rakhiensis Eames, 1952 t;OD(M) Diagnosis: Shell minute, broadly obconic; spire flat; ap- erture narrow; lip strongly thickened, produced on pos- terior corner, smooth, lacking denticulation; strong ex- ternal varix present; strong posterior parietal callus de- posit present near posterior commissure of lip, forming weak posterior notch; siphonal notch absent; columella with 4 strong plications occupying slightly more than hall the aperture. Fossil Record: Eocene of Pakistan. Nomenclature, Synonymy: Currently considered a monotypic genus. Remarks: The 4 columellar plications, smooth, strongly thickened lip that is produced or alate on the posterior corner, presence of an external varix, and absence of a siphonal notch all indicate placement in the Austrogi- nellini and an ancestral relationship to Alaginella. Due to the early occurrence of this group in the Eocene of Pakistan, and the presence of a strong posterior parietal callus deposit, we consider this a separate, valid genus. Genus Alaginella Laseron, 1957 (figure 58) Alaginella La.senm, 1957:286 Carinaginclla Laseron, 1957:286 [TS: Marginella carinata E. A. Smith, 1891; OD (M)] t Cassoginella Laseron, 1957:287 [TS: Marginella palla (Cotton, 1949 t; OD (M)] Triginella Laseron, 1957:280-281 [TS: Marginella malina Hedley, 1915; OD (M)] Type species: Marginella oehracea Angas, 1871; OD (figure 58) Diagnosis: Shell minute to medium, white, biconic, ob- ovate, or subpyriform, usually strongly narrowed ante- riorly; spire low to medium height; aperture narrow to moderately narrow; lip moilerately thickened, denticu- late to smooth, alate posteriorly; external varix present; posterior notch weak or absent; siphonal notch absent; columella with 4 strong unexcavated plications occu- pying slightly more than half the aperture, lacking "false 5th" plication. Type 2 animal; siphon long; mantle pa- pillo.se, extending over external shell surface. G. A. Coovert and H. K. Coovert, 1995 Page 85 Description: Shell (figure 58) minute to medium in size (adult length 1.7-13.0 mm). Color white; surface smooth, glossy. Shape biconic, obovate to subpyriform, usually strongly narrowed anteriorly; weakly to strongly shoul- dered, rarely strongly carinate. Spire usually medium height, rarely very low to low. Aperture narrow to mod- erately narrow, rarely broad, narrowest medially in some species. Lip moderately thickened, thickest medially to posteromedially, posterior corner strongly alate, dentic- ulate to smooth, with a distinct external varix. Shell with- out siphonal notch, posterior notch weak, poorly defined to absent. Shell without obvious ventral callusing. Col- umella with 4 strong continuous plications occupying slightly more than half the aperture length. Internal whorls unmodified. External anatomy: 2 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle covering most or all of external shell surface, papillose with conspicuous, branched papillae; foot nearly as wide as shell, twice as long; animal white, dotted with brown. [Based on de- scription of A. geminata (Hedley, 1912) in Cotton (1944: 16) (as weedingi Cotton, 1944) plus description of A. ochracea (Angas, 1871), ex. unpublished mss., ex. R. Burn and D. R. Hewish]. Internal anatomy: Unknown. Radula: 9 species studied. T\pe 5, uniserial, ribbon short, broad, of 20-55 plates. Rachidian plates usually overlapping, moderately broad (0.019-0.040 mm wide), weakly arched, with 1 1-20 strong cusps on posterior edge. The central cusp usually strongest. Anterior edge of ra- chidian plate slightly to strongly concave, resulting in rectangular to chevron-shaped plates. Shell length: rad- ular width ratio = 86-174. Radular Index = 1.5-4.4. Distribution and Habitat: S. Australian (8 species), Indo- Pacific (8 species). South African (9 species). Intertidal to 1,650 m. Fossil Record: Recent. Miocene and Pliocene of Australia, to Nomenclature, Synonymy: The types of M. ochracea, type species of Alaginella, and M. carinata, type species of Carinaginella. were figured by Kaicher (1981:#2618, #2682). The radulae of these two species, plus M. mal- ina, type species of Triginella, have been studied (pers. obs. of SEMs, ex. Dean Hewish), and all three species are here considered to be congeneric. Conchological dif- ferences are considered to be of specific value only, in- cluding the carinate shoulders of A. carinata. M. palla, the type species of Cassoginella, has an axially costate shell. As in other austroginelline genera, this is considered to be of specific value only. The alate posterior end of lip and lack of a siphonal notch place this species in Alaginella. The principle of first reviser (ICZN Art. 24) is herein employed in choosing Alaginella over the other, more restricted names. Remarks: The alate, flared shoulder of the aperture and the anteriorly strongly narrowed shell, combined with the lack of a strong posterior notch and lack of a "false 5th" columellar plication will serve to distinguish this group. The presence or absence of labial denticulation and axial costae are considered to be of specific difference only. The branched mantle papillae may ultimately prove to be a diagnostic character of this genus. The report on the radula in the original description of Triginella mal- inoides Gabriel, 1962, fig. 4 is apparently erroneous. We have examined SEMs of a radula (ex. Dean Hewish) from a shell intermediate between A. malina (Hedley, 1915) and T. malinoides and it is a typical Alaginella radula. In fact, we consider these two species synonymous. The radula figured by Gabriel is clearly not a marginellid radula and is obviouslv in error. t Genus Hiivia Marwick, 1931 Marginella, subg. Hiwia Marwick, 1931:129 Type species: Marginella (Hiwia) amplificata Mar- wick, 1931 t; OD (M) Diagnosis: Shell small, broadly biconic, strongly nar- rowed anteriorly; spire medium height; strong axial cos- tae present; shoulders angulate to carinate; aperture nar- row; lip thickened, smooth, not denticulate; posterior corner of lip sharply angulate; external varix present; siphonal notch absent; columella with 4 plications oc- cupying more than half the aperture. Fossil Record: Eocene of Australia, to Oligocene of New Zealand. Nomenclature, Synonymy: This group was considered to be a valid genus by Coan (1965:189). Remarks: The two included species (type species and M. aldingae Tate, 1878) appear to be closely related, but otherwise form a distinct, valid genus, apparently extinct since the Oligocene. A relationship to the "Anstroginella Group" and especially Anstroginella is indicated by the 4 columellar plications occupying more than half the aperture, but this group lacks a siphonal notch and has a narrow aperture. Genus Anstroginella Laseron, 1957 (figures 25, 59, 80) Anstroginella Laseron, 1957:285 Plicaginclla Laseron, 1957:285 [TS: Marginella formicula La- marck, 1822; OD] Type species: Marginella muscaria Lamarck, 1822; OD (figure 59) Diagnosis: Shell medium to moderately large, white, yellowish-white, or pale yellowish-orange, biconic, ob- ovate, or subpyriform, usually strongly narrowed ante- riorly; spire low to medium height with shouldered whorls; aperture broad; lip smooth, rapidly thickening posteri- orly with sharpl) defined posterior notch; external varix Page 86 THE NAUTILUS, Vol. 109, Nos. 2 & 3 present; siphonal notch present; heavy ventral callusing usually present; columella with 4 strong plications oc- cupying more than half the aperture. Type 2 animal; siphon long to very long; mantle smooth or weakly pus- tulose, extending over external shell surface. Description: Shell (figure 59) medium to moderately large (adult length 6.2-16.3 mm). Color white, yellowish- white, or pale yellowish-orange; surface smooth, glossy, some species with weak to distinct axial costae. Shape biconic, narrowly to broadly obovate, or subpyriform, usually strongly narrowed anteriorly; weakly to strongly shouldered. Spire low to medium height; spire whorls convex or shouldered, giving uneven profile. Aperture broad, especially anteriorly. Lip moderately to strongly thickened, narrow anteriorly, rapidly thickening poste- riorly but slightly narrowed at posterior fourth, smooth, lacking labial denticulation, with distinct external varix. Shell with distinct siphonal notch, and strong, grooved posterior notch at the junction of body whorl and pos- terior commissure of lip. Shell usually with heavy ventral callusing. Columella with 4 strong continuous plications occupying slightly more than half the aperture length. Internal whorls unmodified. External anatomy: 4 species studied. Type 2 animal; tentacles long, slender; siphon long to very long; mantle smooth or weakly pustulose, covering most or all of ex- ternal shell surface; foot very broad, 2-3 X as wide as shell, about 2 X as long; animal spotted or blotched with white, dark brown, orange, yellow, red, or green. See summary in Ponder and Taylor (1992). Internal anatomy (figure 80): 2 species studied. Same as for "Austroginella Group." Radula (figure 25): 5 species studied. Type 5, uniserial, ribbon short, broad, of 29-61 plates. Rachidian plates overlapping, moderately broad (0.029-0.114 mm wide), weakly arched, with 13-20 strong cusps on posterior edge. Central cusp strongest. Anterior edge of rachidian plate slightly to moderately concave, resulting in chevron- shaped plates. Shell length: radular width ratio = 118- 162. Radular Index = 1.7-3.5. Distribution and Habitat: S. Australian (6 species). In- tertidal to 27 m. Fossil Record: Miocene to Pleistocene of Australia, to Recent. Nomenclature, Synonymy: The nomenclature of these two genus-group names and their type species were dis- cussed in Coovert (1988b), in which the presence of axial costae are considered to be of specific value only. Remarks: This well-characterized group is distin- guished by the presence of 4 strong cohiniellar plications occupying more than half the aperture, combined with the presence of a siphonal and a posterior notch, along with the distinctive lip shape. For an account of pred- atorv shell drilling and anatomv, see Ponder and Tavlor (1992). Genus Mesoginella Laseron, 1957 (figure 60) Mesoginella Laseron, 1957:282 Deviginella Laseron, 1957:283-284 [TS: Marginclla brachia Watson, 1886, = Marginella {Glabella} brachia Watson, 1886; OD] t Hianoginetla Laseron, 1957:288 [TS: Marginella physa Cot- ton, 1949 t: OD (M)] Sinuginella Laseron, 1957:282 [TS: Marginella inconspicua G. B. Sowerby II, 1846; OD] Spiroginella Laseron, 1957283 [TS: Marginella leia Cotton, 1944, = M. turbinata G B. Sowerby 11, 1846; OD (M)] t Urniginella Laseron, 1957:287 [TS: Marginella cassidijormis Tate, 1878 t; OD (M)] Type species: Marginella turbinata G. B. Sowerby II, 1846; OD (M) (figure 60) Diagnosis: Shell small to medium, white or yellowish- white, rarely brownish-orange or with pale bands, bi- conic, obconic, obovate, or broadly cylindrical, not strongly narrowed anteriorly; spire low to medium height with evenly contoured whorls; aperture moderately nar- row; lip smooth to denticulate, moderately to strongly thickened, thickest medially; external varix present; weak siphonal notch usually present; posterior notch weak to absent; ventral callusing usually absent; columella with 4 strong plications occupying slightly less to slightly more than half the aperture, 4th plication often remote. Tvpe 2 animal; siphon long; mantle smooth or weakly pustu- lose, extending over external shell surface. Description: Shell (figure 60) small to medium in size (adult length 2.5-11.0 mm). Color usually white to yel- lowish-white, semi-opaque to translucent, rarely brown- ish-orange or with pale yellow bands; surface smooth, glossy, some species with weak to distinct axial costae. Shape narrowly to broadly obovate, obconic, biconic, or broadly cylindrical, not strongly narrowed anteriorly; weakly to strongly shouldered. Spire low to medium height with evenly contoured whorls. Aperture moder- ately narrow, usually wider anteriorly. Lip moderately to strongly thickened, thickest medially, usually smooth, some species with weak to moderately strong labial den- ticulation, with distinct external varix. Siphonal notch nearly absent to weak, rarely strong, distinct or com- pletely absent. Posterior notch weak, poorly defined or absent. Shell usually without evident ventral callusing. Columella with 4 strong continuous plications, 4th often remote, rarely with parietal tubercle appearing as "false 5th ' plication. Plications occupying slightly less to slight- ly more than half the aperture length. Internal whorls unmodified. External anatomy: 5 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle smooth or weakly pustulose, asymmetrically extending over exter- nal shell surface; foot narrow to broad, slightly narrower to 1 '/2 .\ as wide as shell, about I'/s X as long; animal variously spotted, blotched, or with lines of various colors. G. A. Coovert and H. K. Coovert, 1995 Page 87 including white, yellow, orange, brown, and black. See summary in Coovert (1987g). Internal anatomy: 1 species studied. Same as for "Aus- trogiiiella Group. Radula: 16 species studied. Type 5, uniserial, ribbon short, broad, of 19-75 plates. Rachidian plates usually overlapping, moderately broad (0.018-0.050 mm wide), weakly arched, with 9-22 strong cusps along posterior edge. The central cusp (or two subcentrals) is the strong- est. Anterior edge of rachidian plate slightly to strongly concave, resulting in rectangular to chevron-shaped plates. Shell length: radular width ratio = 101-221. Radular Index = 1 5-7.1. Distribution and Habitat: Neozelanic (13 species), S Australian (13 species), Indo- Pacific (7 species). Intertidal to 640 m. Fossil Record: Miocene, Pliocene, and Pleistocene of Australia, to Recent. INomenclature, Synonymy: The nomenclature of Me- soginella and Sinuginella and their type species were discussed by Coovert (1988b). The type of M. inconspi- cua was figured by Kaicher (1981;#2702). M. brachia, type species of Deviginella, falls well within the limits of Mesoginella. The principle of first reviser (ICZN Art. 24) is here employed in choosing the better known Me- soginella over Deviginella. The holotype of M. leia was figured in Hewish and Gowlett-Holmes (1991;64, figs. E-F) and synonymized with M. turbinata, thus placing Spiroginella in synonymy. The two fossil genera Hian- oginella and Urniginella are here synonymized, based on conchological characters falling within the limits of Mesoginella. Laseron (1957:284) used several nomina nuda that are apparently manuscript names for genera synonymized here. These should not be further consid- ered. Remarks: The presence or absence of axial costae are not considered to be a genus-level character in this group. The 4th columellar plication is often remote, i.e. sepa- rated farther from the other 3 plications. This is a rather diverse genus but all species possess the same type of radula and conchological characters. Genus Closia Gray, 1857 (figure 61) Closia Gray. 1857:36 Type species: Closia sarda (Kiener, 1834), = Margi- nella sarda Kiener, 1834; M (figure 61) Diagnosis: Shell medium to moderately large, white or colored, obovate; spire immersed; lip thickened, dentic- ulate; external varix present; siphonal notch and posterior notch absent; columella with 4 thin, sharp plications crowded anteriorly. Description: Shell (figure 61) medium to moderately large (adult length 9.9-48.0 mm). Color grayish-white, without other markings; pale rose, pink, or orangish- yellow, with lip and apical spot darker brownish-rose or yellowish-orange; or pale yellowish- to light orange with paler mid-body band and lip darker with tiny white specks. Shell surface smooth, glossy. Shape obovate, weakly to strongly narrowed anteriorly, with rounded to angulate shoulders. Spire immersed. Aperture moder- ately narrow, wider anteriorly. Lip moderately to strong- ly thickened, strongly to very strongly denticulate in adults, with a distinct external varix. Shell without si- phonal notch or posterior notch. Shell usually with evi- dent ventral callus deposits anteriorly and especially pos- teriorly. Some with heavy parietal callus wash abruptly ending at entrance to aperture, creating appearance of collabral parietal callus ridge. Columella with 4 thin, sharp, continuous plications, crowded anteriorly, occu- pying slightly less than Va to Vi aperture length. Internal whorls presumed unmodified. External anatomy: 1 species studied. Animal white, semi-transparent. [From description of C. limpida Boz- zetti, 1992:11, = C. majuscula (Martens, 1880)]. Pre- sumed Type 2 animal. Internal anatomy: Unknown. Radula: Unknown. Distribution and Habitat: Indo-Pacific (3 species), re- stricted to western Indian Ocean. Recorded from 20 to 140 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: Coan (1965:189), followed by others, considered this group to be a subgenus of Bullata, apparently based on the similarity to B. lilacina (G. B Sowerby II, 1846). Bullata, which is included in Prunini based on its radula, anatomy, and fossil ancestry, has a tendency toward heavy ventral callusing and stronger columellar plications that are not nearly as crowded anteriorly as in Closia. Although the radula and animal are unknown, we feel that Closia is not closely related to Bullata and has a different ancestry. Remarks: Closia has been variously associated with Bul- lata and Ovaginella. Conchologically, it comes closest to Ovaginella, with which it shares shell shape, and thin, sharp, anteriorly crowded columellar plications. The coarser labial denticulation, distinct shell pigmentation, heavier posteroventral callus deposit, and Indian Ocean distribution serve to distinguish it. Until the radula and animal are known, these two groups are provisionally considered distinct. Genus Ovaginella Laseron, 1957 (figure 62) Ovaginella Laseron, 1957:280 Type species: Marginella ovulum G. B. Sowerby II, 1846; OD (figure 62) Page 88 THE NAUTILUS. Vol. 109, Nos. 2 & 3 69 FiKures 61-69. Shells of tvpe .species of marginellid genera, ventral views. 61. Closia ^ania (Kiener, l*^*^-!* GAG M1341 off Anibanja, N.E. Madagascar.' dredged. Length 17.6 mm. 62. Ovaginella ovulum (G. B. Sowerby II, 1846). Type, BM(NH) 80.9.^5, locality unknown. After photograph in Kaicher (1981:#2647). Length ca. 9 mm. 63. Balanetta baylei ^"'f'^^'^'^lJ^'^ ^-.^^ M1607 Margaret River, Western Australia, on beach. Length 10.5 mm. 64. Voharina mitrella (Risso, 1826). OAC MZbM, balina Bav Malta, dredged on sandy bottom at 6 m. Length 9.3 mm 65. Prunum prurium (Cimelin, 1791). GAG M1393, Venezuela^ Length 26 9 mm. 66. Bullata Imllaia iBorn, 1778). GAC: M1787, Praia de Forte Gragoata, Niteroi, Rio de Janeiro, Brazil. Length G. A. Coovert and H. K. Coovert, 1995 Page 89 Diagnosis: Shell small to medium, white, elliptic to ob- ovate; spire immersed; lip thickened, usually smooth, rarely denticulate; external varix present; siphonal notch and posterior notch absent; columella with 4 anteriorly crowded plications. Description: Shell (figure 62) small to medium in size (adult length 4.2-10.1 mm). Color translucent to semi- opaque white; surface smooth, glossy. Shape narrowly to broadly elliptic to obovate, weakly to strongly shoul- dered. Spire immersed. Aperture moderately narrow, slightly wider anteriorly. Lip moderately thickened, usu- ally smooth, rarely weakly denticulate, with distinct ex- ternal varix. Siphonal notch weak or absent; posterior notch absent. Shell usually without evident ventral cal- lusing. Columella with 4 continuous plications crowded anteriorly, occupying distinctly less than half aperture length. Internal whorls unmodified. External anatomy: Unknown. Internal anatomy: Unknown. Radula: 2 species studied. Type 5, uniserial, ribbon short, broad, of 20-31 plates. Rachidian plates overlap- ping, moderately broad (0.025-0.035 mm wide), weakly arched, with 8-14 strong cusps along posterior edge. The central cusp is the strongest. Anterior edge of rachidian plate slightly concave, resulting in rectangular to chev- ron-shaped plates. Shell length: radular width ratio = 170-285. Radular Index = 1.8-3.5. Distribution and Habitat: Neozelanic (2 species), S Aus- tralian (2 species). Recorded from 3 to 370 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: The type species of Ova- ginella, M. ovulum, is figured by Kaicher (1981:#2674) and herein (figure 62). Coan (1965:189) placed Ovagi- nella as a subgenus of Balanetta. but the latter has only 3 columellar plications and a narrower shape. Remarks: Although Balanetta and Ovaginella both have a Type 5 radula, they are considered distinct based on conchological differences. Genus Balanetta Jousseaume, 1875 (figure 63) Balanetta Jousseaume, 187.5:168, 269 Type species: B. baylei Jousseaume, 1875; M (figure 63) Diagnosis: Shell minute to medium, white, cylindrical to narrowly obovate; spire immersed; lip thickened, smooth; external varix present; with weak or absent si- phonal notch; posterior notch absent; columella with 2 or 3 plications crowded anteriorly. Description: Shell (figure 63) minute to medium in size (adult length 1.5-12.0 mm). Color translucent to opaque white or yellowish-white; surface smooth, glossy. Shape cylindrical to narrowly obovate, weakly to strongly shoul- dered. Spire immersed. Aperture moderately narrow, distinctly wider anteriorly. Lip moderately thickened, smooth, not denticulate, with a distinct external varix. Siphonal notch weak or absent, posterior notch absent. Shell without evident ventral callusing. Columella with 3 continuous plications (one species with 2), crowded anteriorly, occupying less than Vi aperture length. In- ternal whorls presumed unmodified. External anatomy: Unknown Internal anatomy: L'nknown. Radula: 1 species studied. Type 5, uniserial, ribbon relatively short, broad, of 29-37 plates. Rachidian plates overlapping, moderately broad (0.027-0.031 mm wide), weakly arched, with 9-14 strong cusps on posterior edge. Central cusp strongest. Anterior edge of rachidian plate slightly concave, resulting in rectangular to chevron- shaped plates. Shell length: radular width ratio = 185- 239. Radular Index = 2.6-3.1. Distribution and Habitat: S. Australian (3 species), Indo- Pacific (1 species). Intertidal to 183 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: The type species was ded- icated to M. Bayle in the original description by Jous- seaume (1875:269, 274) but spelled "baylii" in the text and "baylei" in the figure caption (op. cit., pi. 8, fig. 5). This is to be considered a case of multiple original spell- ings, which is dealt with in the ICZN [Art. 32(b)(i)]. This section, then, is subject to Section 32(c)(ii), which con- siders an incorrect original spelling one in which the original publication itself shows clear evidence of an inadvertent error. Because Jousseaume intended the spe- cies to be dedicated to M. Bayle, it is clear that the spelling "baylei" is the correct one. Tomlin (1917:252) considered "baylei" an error. Remarks: The immersed spire, lack of labial denticu- lation, and presence of an external varix and only 3 columellar plications, distinguishes this group. Tribe PRUNINI Coovert and Coovert, new tribe Diagnosis: Shell small to very large, white, uniformly colored, patterned, or banded; spire immersed, or low to tall; lip thickened, smooth to denticulate; external varix present or absent; siphonal notch usually absent; posterior notch absent; columella with 2-6 plications occupying half or less of the aperture. Type 2 animal; siphon long 57.6 mm. 67. Rivomarginella morrisoni Brandt, 1968. GAC Ace. # 11-89, Prachin River, Kabinburi. Thailand. Length 10 0 mm. 68. Cryptospira tricincta (Hinds, 1844). GAC M1253, Formosa Strait, S.W. Taiwan, dredged at .37 m. Length 23.8 mm. 69. Hyalina pallida (Linne, 1758). GAC M2068, St. Croix, U.S. Virgin Islands. Length 14.3 mm. Page 90 THE NAUTILUS, Vol. 109, Nos. 2 & 3 to very long; mantle smooth, pustulose, rarely distinctly papillose, usually extending over external shell surface. Type 6 radula or non-radulate. Marginellid buccal pouch present, absent in non-radulate species; odontophoral cartilages present in radulate species, fused anteriorly or both anteriorly and posteriorly; valve of Leiblein absent; esophageal caecum present; gland of Leiblein with long, convoluted duct and a terminal bulb, passing through nerve ring and emptying into anterior end of proboscis; paired salivary glands ascinous or tubular, either con- tained within proboscis or free, ducts either attached to walls of esophagus or free; single accessory salivary gland present or absent, ascinous or tubular. Genus Voharina Hinds, 1844 (figures 38, 64) Marginella, section Volvarina Hinds, 1844:75 Type species: Marginella nitida Hinds, 1844, = Mar- ginella (Volvarina) nitida Hinds, 1844, = Valuta mi- irella Risso, 1826; SD Redfield, 1870:221 (figure 64) Diagnosis: Shell small to moderately large, color trans- lucent white, amber, or brown, usually with darker bands; spire immersed, or low to tall; lip thickened, smooth, not denticulate; external varix usually absent, but weak to strong in some species; siphonal notch absent or weak; columella with 3 or 4 plications, some species with weak- er 5th or 6th, combined occupying half or less of the aperture. Type 2 animal; siphon long to very long; mantle smooth or usually pustulose, rarely distinctly papillose, usually extending over external shell surface. Type 6 radula. Description: Shell (figure 64) small to moderately large (adult length 2.6-22.4 mm). Color translucent to opaque white, amber, or brown, usually with yellow, orange, brown, or reddish spiral bands, rarely axially streaked; surface smooth, glossy. Shape elongate to moderately broadly cylindrical, elliptic, narrowly to broadly obovate, or oblong, rarely elliptic-cylindrical with immersed spire; weakly to strongly shouldered. Spire rarely immersed, usually low, medium, or tall. Aperture narrow to mod- erately broad, rarely broader, usually wider anteriorly. Lip narrowly to strongly thickened, smooth, not dentic- ulate, usually lacking an external varix, some species with weak to strong external varix. Siphonal notch rarely weak, usually absent; posterior notch absent. Ventral callusing usually not evident, present in one species group. Col- umella usually with 4 continuous plications; rarely with 3 plications and "false 4th" plication that joins short collabral parietal callus ridge; or 5 plications, the 5th not continuous past V2 whorl internally, plus often "false 6th" plication or parietal lira posteriorly. Plications usually occupying less than half, but some species up to half the aperture length. Internal whorls umnodified. External anaton- rad, 1868 t; M] Egouena Jonsscaume, 1875: 167, 192 |TS: E rgoiicn Jous- seauine, 1875, = Marginella amygdala Kiener, 1841; T] G. A. Coovert and H. K. Coovert, 1995 Page 91 t PorccUanella Tryon, 1882; 16, (non White in MacGillivray, 1852) [TS; P. hella Conrad, 1868, = Prunum bclla C:onrad, 1868 1; OD (M)][publ as nomeii mtdiiin in Conrad, 1863: 564, TS: P. hella Conrad, 1863, nomen nudum, M] Marginclla. subg. \'olvarina, section Lcptcgouana Woodring, 1928:237-238 [TS: Valuta guttata Dillwyn, 1817, OD] Type species: Valuta prunum Gnielin, 1791; M (figure 65) Diagnosis: Shell small to large, usually thick, opaque, white or uniformly colored, often patterned, but usually not simply with dark, narrow bands; shell variously shaped; lip moderately to strongly thickened, smooth to denticulate; external varix usually present; siphonal notch usually absent; posterior notch absent; shell usually with heavy ventral callusing; columella with 4 plications oc- cupying half or less of the aperture. Type 2 animal; siphon long to very long; mantle smooth, extending over external shell surface. Type 6 radula. Descriplion: Shell (figures 1, 65) small to large (adult length 3.3-44.1 mm). Shell usually thick, opaque, rarely translucent, white or colored tan, gray, brownish-gray, orange, or pink; uniformly colored, streaked, or spotted, or with paler spiral bands on colored species, some species with opaque white flecks overlaying other patterns, rare- ly banded with darker color. Surface smooth, glossy. Shape obovate, oblong, subtriangular, biconic, or rarely sub- cylindrical; weakly to strongly shouldered. Spire rarely immersed, usually low to medium height. Aperture nar- row to moderately broad, wider anteriorly. Lip moder- ately to very strongly thickened, in one group much thinner anteriorly, smooth, often weakly to distinctly denticulate; usually with very distinct, rarely duplicate, external varix that is weak to absent in some species. Siphonal notch usually absent, rarely weakly developed; posterior notch absent. Ventral callus usually heavy, of- ten with strongly produced deposit near posterior com- missure of lip. Columella usually with 4 continuous pli- cations, rarely with weak 5th plication. Plications usually occupying half or less of aperture length. Internal whorls unmodified. External anatomy (figures 7, 8): 17 species studied. Type 2 animal; tentacles long, slender; siphon long to very long; mantle smooth, symmetrically extending over ex- ternal shell surface, often nearly completely covering shell; foot broad and long, about l'/2 X shell width, 2 X shell length; animal variously marked with tiny dots, spots, blotches, or lines of various colors. Internal anatomy (figures 31, 39-42): 6 species studied. As in tribe. Marginellid buccal pouch present; odon- tophoral cartilages fused anteriorly or both anteriorly and posteriorly; paired salivary glands ascinous or tubular, free, ducts either attached to walls of esophagus or free; single tubular accessory salivary gland present or absent. Radula (figures 17-20, 27): 22 species studied. Type 6, uniserial, ribbon short, broad, of 24-59 plates. Rachidian plates overlapping, broad (0.096-0.920 mm wide), nearly flat, with 16-45 sharp cusps on posterior edge. Anterior edge of rachidian plate generally straight, producing elongate, rectangular "comb-like" plates. Shell length: radular width ratio = 30-103. Radular Index = 0.6-2.9. Distribution and Habitat: Indo-Pacific (2 species), E. Pacific (7 species), W. Atlantic (52 species), W. African (3 species). Intertidal to 1,840 m. Fossil Record: Eocene of Mississippi, Oligocene to Pleistocene of W. Atlantic, to Recent. Nomenclature, Synonymy: The four genus-group names synonymized under Prunum differ mainly in degree of callusing, spire development, and degree of thickening and denticulation of the outer lip, all considered to be of specific value only. Egouena is in the same species group as P. prunum based on its type species E. egouen, a junior synonym of M. amygdala. This is supported by our observations of specimens of P. marginatum (Born, 1778), in which subadults are extremely similar to P. prunum, whereas fully adult specimens with heavy callus deposits are obviously close to P. amygdalum. All share the same brown apertural coloration and combined with several other species form a closely related species group. The 3 multiple original spellings of Egouena were listed and discussed in Coan (1965:189), where Neave (1939: 2:199) was listed as the first revisor. The other 3 genus- group names are not quite as closely related, but certainly appear to be congeneric. The holotype of Volutella ovi- formis was figured by Gardner (1937:pl. 47, figs. 11-12). The type of P. helium was figured by Olsson and Har- bison (1953: pi. 30, fig. 4). Coan (1965:189) synonymized Egouena, Porcellanella, and Lcptegouana under Prun- um. Roth (1978:8) tentatively considered Egouena and Leptegouana synonyms of Microspira. Some of these genus-group names could be employed as subgeneric groups after all Prunum and Volvarina are fully revised, but this would probably necessitate introducing addi- tional formal names. We feel that recognition of informal species groups would be a better solution. Much research remains to be done along these lines, especially with the anatomy and fossil fauna. Remarks: The concept of this genus outlined in Coovert (1988c) has been further modified, partially based on the work of Gofas (1989b) and Gofas and Fernandes (1992) on Volvarina. These authors placed the majority of West African species, formerly included in Prunum, in Vol- varina. They stated that the separation of these two gen- era was largely subjective as far as shell features, a con- tention with which we agree. Species of Prunum have a smooth mantle, whereas the West African Volvarina have a pustulose or papillose mantle. The group of non- varicose, narrow Volvarina often have a smooth mantle, but in at least one species group {"rubella group, see Coovert & Coovert, 1990), their radula distinguish them from Prunum. Also, the ontogenetic development of the thickened outer lip in varicose and non-varicose species may further distinguish these genera (cf. section on shell Page 92 THE NAUTILUS, Vol. 109, Nos. 2 & 3 morphology). The key to genera will serve to identify these two groups based on current concepts. A satisfactory solution to this complex situation will only be realized through a complete revision of all species groups in both genera, based on shell and anatomical features in combination with a study of the fossil record. We feel that the large group of western Atlantic species assigned to Prunum, both varicose and non-varicose, forms a natural assemblage distinct from the western Atlantic species assigned to Volvarina. Three major spe- cies radiations apparently occurred: eastern Atlantic Vol- varina with a pustulose mantle; western Atlantic Prunum with a smooth mantle; and a much more widespread group of narrow, non-varicose Volvarina. Genus Bullata Jousseaume, 1875 (figure 66) Bullata Jou.sseaume, 1875:167, 250 Marginclla, subg. Volutella Swaiiison, 1830:(2)1, Marginella pi. 1 (non Perry, 1810) [TS: Marginella bxdlata Lamarck, 1822, = Voluta Inillata Born, 1778; OD] Gibbertilina Monterosato, 1884:139 [invalid emendation, as "nom sost ' ] Type species: Bullata bullata (Born, 1778), = Voluta bullata Born, 1778; T (hgure 66) Diagnosis: Shell moderately large to very large, colored and banded or patterned, elliptic to oblong or obovate; spire immersed or nearly so; lip thickened, denticulate in adults; external varix present; siphonal notch absent; columella with 4 strong plications occupying less than half the aperture but are not crowded anteriorly. Type 2 animal; siphon long; mantle extension undetermined. Type 6 radula. Description: Shell (figure 66) moderately large to very large (adult length 16.8-90.0 mm). Color yellowish-or- ange to orangish- or pinkish-brown, spirally banded or with white spots, lip pink, yellow, or orange, darker than shell color. Shell surface smooth, glossy. Shape elliptic to oblong or obovate, moderately to strongly shouldered. Spire immersed or nearly so. Aperture narrow to mod- erately broad, wider anteriorly. Lip moderately to strongly thickened, weakly to strongly denticulate in adults, with a distinct external varix. Siphonal notch, posterior notch absent. Parietal callusing weakly to strongly developed, especialK posteriorly, absent in type species. Columella with 4 continuous plications occupying less than half the aperture length. Internal whorls unmodified. External anatomy: 1 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle extension undetermined; foot moderately broad, about 1 '/s X shell width, slightly longer than shell; animal marked with fine, narrow red lines on cream background. [Based on dissection of Rnllata bullata]. Internal anatomy: 1 species studied. As in tribe. Mar- ginellid buccal pouch present. Radula: 1 species studied. Type 6, uniserial, ribbon short, broad, of 47 plates. Rachidian plates overlapping, broad (0.825 mm wide), nearly flat, with 40 sharp cusps along posterior edge. Anterior edge of rachidian plate generally straight, resulting in elongate, rectangular "comb-like" plates. Shell length: radular width ratio = 63. Radular Index = 1.2. Distribution and Habitat: W. Atlantic (4 species). Re- corded from 1 to 60 m. Fossil Record: Miocene and Pliocene of W. Atlantic, to Recent. Nomenclature, Synonymy: Nomenclature of Gibberti- lina was discussed in Coovert (1987h:27). Remarks: The large, patterned shells with an immersed spire and 4 moderately heavy columellar plications not crowded anteriorly serve to distinguish this group. The presence of an esophageal caecum clearly places this genus in the Prunini. As here defined, this group is re- stricted to the Caribbean province, where it evolved in the Miocene as a direct offshoot of Prunum. Genus Rivomarginella Brandt, 1968 (figure 67) Rivomarginella Brandt, 1968:275 Type species: R. morrisoni Brandt, 1968; (OD) M (fig- ure 67) Diagnosis: Shell small to medium, translucent, lightly colored, shape obconic to subpyriform; spire medium height; aperture broad; lip thickened, smooth; external varix present; siphonal notch absent; with minutely gran- ulated apical and ventral callusing; columella with 4 plications occupying half the aperture length. Type 2 animal; siphon long; mantle smooth, extending over ex- ternal shell surface. Type 6 radula. Description: Shell (figure 67) small to medium in size (adult length 5.4-11.3 mm). Color translucent >ellowish- white to brownish-white, some species with 1 or 2 obscure spiral bands; surface smooth, glossy. Shape broadly ob- conic to subpyriform, moderately to strongly shouldered. Spire medium height. Aperture moderately broad to broad, wider anteriorly. Lip moderately thickened, smooth, lacking denticulation, with a distinct external varix. Siphonal notch, posterior notch absent. Shell with minutely granulated ventral and apical callusing. (Col- umella with 4 continuous plications occupying half the aperture length. Internal whorls presumed unmodified. External anatomy: 2 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle smooth, ex- tending almost completely over external shell surface; loot moderately broad; animal pigmented with hlackish- browii, black, or yellow dots or blotches. The penis was described by Brandt (1968:277) as long, simple, and lack- ing appendages. G. A. Coovert and H. K. Coovert, 1995 Page 93 Internal anatomy: Ihiknown. Radula: 2 species studied. Type 6, uiiiserial, ribbon short, broad, of 38-47 plates. Rachidian plates overlap- ping, broad, nearly flat, with 18-29 sharp cusps along posterior edge. Anterior edge of rachidian plate generally straight, resulting in elongate, rectangular "comb-like' plates. No further ilata available. Distribution and Habitat: Indo-Pacific (2 species), found in freshwater in S.E. Asia. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: See Brandt (1968:pl. 10, fig. 63) for a photograph of the holotype of the type species. Remarks: Subadult shells show that the lip is first re- flected, then gradually thickened by callus deposits. This is unlike Volvarina, which initially has the lip curved in. Thus, we consider Rivomarginella to be more closely allied to the Prunum lineage. Eraloidea was considered by Brandt (1968:275) to be the closest relative, but it is apparently non-radulate and has stronger columellar pli- cations occupying more than half the aperture. For fur- ther information see Brandt (1974), Coomans and Clover (1972), and Djajasasmita and Coomans (1980). Genus Cryptospira Hinds, 1844 (figure 68) Marginella. section Cryptospira Hinds, 184476 Type species: Marginella tricincfa Hinds, 1844, = Mar- ginella (Cryptospira) trieincta Hinds, 1844; SD (M) Gray, 1847:142 (figure 68) Diagnosis: Shell medium to large, opaque, uniformly colored or with bands or patterned; spire immersed or low; lip thickened, smooth or denticulate; external varix present; siphonal notch absent; columella with 4 to 6 plications. Type 2 animal; siphon long; mantle smooth, extending over external shell surface. Type 6 radula. Description: Shell (figure 68) medium to large (adult length 6.2-46.0 mm). Color usually uniformly opaque gray or flesh-colored to pale brown, many with narrow spiral lines, undulating longitudinal zig-zag lines, lon- gitudinal streaks, or spiral lines crossed by longitudinal streaks, lip often differentially colored. Often relatively thick-shelled, surface smooth, glossy. Shape cylindrical, elongate to broadly elliptic, obovate, pyriform, or sub- triangular, moderately to strongly shouldered. Spire im- mersed or low. Aperture narrow to moderately broad, wider anteriorly. Lip moderately to strongly thickened, smooth or with weak to strong denticulation in adults, with a distinct external varix that is rarely duplicate. Siphonal notch absent or at most very weakly developed; posterior notch absent, weak notch present between pos- terior parietal callus deposit and posterior lip commis- sure. Shell often with anterior or posterior ventral parietal callusing. Columella with 4 to 6 continuous plications occupying distinctly less than half to more than half the aperture length. Internal whorls unmodified. External anatomy: 4 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle smooth, ex- tending symmetrically or nearly so over external shell surface; foot moderately broad, about l'/2 X shell width, 1 '/2 X length; animal uniformly colored or brightly marked with streaks of red on yellow ground color. Internal anatomy: Unknown. Radula: 4 species studied. Type 6, uniserial, ribbon short, broad, of 36-50 plates. Rachidian plates overlap- ping, broad (0.158-0.403 mm wide), nearly fla', with 9- 28 sharp cusps along posterior edge. Anterior edge of rachidian plate generally straight, resulting in elongate, rectangular "'comb-like" plates. Shell length: radular width ratio = 35-77. Radular Index = 1.4-2.3. Distribution and Habitat: Indo-Pacific (15 species). Re- corded from 0.6 to 123 m. Fossil Record: Pliocene of Java, to Recent. Nomenclature, Synonymy: The earliest type species designation is by Gray (1847:142), as noted in Coan (1965: 189). Besides being predated by Gray, Cossmann's (1899: 94) designation of Marginella quinqueplicata Lamarck, 1822, as type species, is invalid because this was not an included species of Hinds. See Palmer (1937:418) for comments on nomenclature, but note that Hinds' useage of division places this taxon as a genus-group name (ICZN, Art. lOe). We do not consider Euryentome to be closely related to Cryptospira. Coan (1965:189) placed Cryp- tospira as a subgenus of Bullata. We consider these two groups to have separate origins, Bullata being a direct descendant of Caribbean Prunum. whereas Cryptospira is restricted to the western Indo-Pacific where it evolved. Remarks: See Coomans (1969) for further information. The large, thick, colored, frequently patterned shells, often with 5 or 6 columellar plications, serve to differ- entiate this group. One species group within the genus has distinctively gray-colored shells. As defined and re- stricted geographically, this genus forms a compact, closely related group. Genus Hyalina Schumacher, 1817 (figures 32, 69) Hyalina Schumacher, 1817:234 Marginella, subg Volvarina. section Neovolvaria Fischer, 1883: 602 [TS: Marginella pallida (Linne, 1767), = Bulla pallida Linne, 17.58; M] Type species: Hyalina pellucida Schumacher, 1817, = Bulla pallida Linne, 1758; M (figure 69) Diagnosis: Shell small to large, white to lightly colored, hyaline or translucent, thin-shelled; spire low to medium height; aperture broad; lip thin, smooth; external varix weak to absent; siphonal notch absent; ventral callusing Page 94 THE NAUTILUS, Vol. 109, Nos. 2 & 3 absent; columella with 2 to 4 plications occupying less than half the aperture. Type 2 animal; siphon long; man- tle smooth. Non-radulate. Description: Shell (figure 69) small to large (adult length 4.0-29.0 mm). Color white, hyaline or translucent, cream to amber or brown, rarely with faint yellowish-white bands or numerous spiral lines. Thin-shelled, surface smooth, glossy. Shape broadly cy lindric to obovate, weak- ly to moderately shouldered. Spire low to medium height. Aperture moderately broad to broad, usually distinctly wider anteriorly. Lip moderately thin, smooth, lacking denticulation; e.xternal varix weak to absent. Siphonal notch, posterior notch, ventral callusing absent. Colu- mella with 2-4 continuous plications occupying distinctly less than half the aperture length. Internal whorls un- modified. External anatomy: 3 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle smooth, ap- parently extending over external shell surface; foot mod- erately broad, about l'/2 X shell width, I'/a X length; animal white or translucent, uniformly colored or spotted and flecked with orange and black. Internal anatomy (figure 32): 2 species studied. As in tribe. Marginellid buccal pouch absent, paired salivary glands tubular, free, ducts free, single accessory salivary gland present, tubular. Radula: Non-radulate, completely lacking buccal mass, including odontophore. 3 species known or strongly sus- pected to be non-radulate. Distribution and Habitat: Indo- Pacific (1 species), W. Atlantic (3 species), Magellanic / Antarctic (2 species). South African (5 species). Intertidal to 1,340 m. Fossil Record: Pliocene of Florida, to Recent. Nomenclature, Synonymy: Coan and Roth (1976) gave a complete discussion of Uijalina and its type species. They designated (op. cit. fig. 1) the same specimen as neotype for both Bulla pallida Linne, 1758 and Hyalina pellucida Schumacher, 1817, placing Neovolvaria as an objective synonym of Hyalina. Remarks: The complete lack of a radula and buccal mass at first suggested placement of this genus with the non-radulate marginellines, but conchological features did not conform. Subsequent dissection of a specimen of H. pallida revealed the presence of an esophageal cae- cum, a diagnostic feature of the Prunini. The thin, trans- lucent shells, a weak or absent external varix, and 2, 3, or 4 columellar plications will generally serve to distin- guish this genus. Tribe MARGINP:LL1NI Fleming, 1828:328 Diagnosis: Shell minute to very large, white, uniformly colored, or patterned, biconic, obconic, or broadly obo- vate; spire low to tall; lip thickened, smooth or dentic- ulate; external varix present; siphonal notch present or absent; columella with 4 or 5 strong plications occupying half or more of the aperture. Type 2 animal; siphon long; mantle smooth or papillose, variably extending over ex- ternal shell surface. Non-radulate, lacking entire buccal mass, including odontophore and cartilages. Proboscis rounded, blunt, or distally pointed; marginellid buccal pouch absent; valve of Leiblein absent; esophageal cae- cum absent; gland of Leiblein with a terminal bulb and a long, convoluted duct that passes through nerve ring and empties into anterior end of proboscis; paired sali- vary glands ascinous or tubular, either contained within proboscis or free, ducts either attached to walls of esoph- agus or free; single accessory salivary gland present, tu- bular. t Genus Stazzania Sacco, 1890a Marginella. siibg. Stazzania Sacco, 1890a:138 (245); 1890b:26 (318) Type species: Marginella (Stazzania) emarginata Sis- monda, 1847, = Marginella emarginata Sismonda, 1847 t; M Diagnosis: Shell miimte to moderately large, narrowly to broadly biconic or broadly obovate; spire medium to tall; aperture narrow; lip thickened, smooth or dentic- ulate, often with stronger or single posterior denticle; external varix present; siphonal notch absent; columella with 4 or 5 strong plications occupying more than half the aperture; coUabral parietal callus ridge present, usu- ally connecting with bifurcated outer ends of plications. Fossil Record: Eocene of France, to upper Miocene of Italy. Nomenclature, Synonymy: Many European paleontol- ogists synonymize Dentiinargo under Stazzania (e.g. Gougerot & Le Renard, 1979; Nieulande, 1981; Le Ren- ard & Nieulande, 1985), apparent!} based on similarities in shell shape and the presence of bifurcated columellar plications in some species of both groups. The bifurcated plications in Stazzania result from thin, paired callus ridges on their distal ends joining a collabral parietal callus ridge However, in fossil species we assign to Den- tiinargo, bifurcated columellar plications are fundamen- talK different, being thick and broad, with a U-shaped notch distally that creates the bifurcate appearance. Spe- cies of Dentimargo with a collabral callus ridge do not have this ridge intersecting the distal ends of the plica- tions. Most Recent species of Dentimargo have simple, unmodified plications. The t\pe species of Stazzania is significantly larger than the relatively small Eocene spe- cies herein assigned to the same group, but has a very similar shape. Remarks: Stazzania is here considered to be an extinct, separate lineage from the contemporaneous Dentimar- go. Dentimargo. besides differing in plications, is gen- erally narrower with a broader aperture. G. A. Coovert and H. K. Coovert, 1995 Page 95 Figures 70-73. Shells of type species of margineiiid genera, ventral views. 70. Dcntiniargo dentifcra (Lamarck, 1803). GAC Ace. # 35-iS7, fossil, Lutetian of the Eocene, Paris Basin, Fercourt, France. Length 4.6 mm. 7L Eratoidca margarila (Kiener, 1834). GAC M2079, Water Island, St. Thomas, Virgin Islands. Length 8.0 mm. 72.^Margim'lla glabella (Linne, 1758). GAC M1322, Cape Verde Islands. Length 38.7 mm. 73. Glabella faba (Linne, 1758). GAC M1720, Dakar, Senegal, West Africa, under rocks at 15 m. Length 18.5 mm. Genus Dentimargo Cossmann, 1899 (figures 33, 70) Marginella. subg Marginella, section Dentimargo (.'.ossTiMinn, 1899:90 Volvannella Habe, 1951:101-102 [TS: V. makiyamai Habe, 1951; OD] Marginella, subg. Elmrnospira Olsson & Harbison, 1953:201- 202 [TS: Marginella elmrneola Conrad, 1834 f; OD] Longinella Laseron, 1957:286, (non Gros & Lestage, 1927) [TS: Marginella maugeana Hedley, 1915; OD] Type species: Marginella dentifera Lamarck, 1803 t; OD (figure 70) Diagnosis: Shell small to medium, white or uniformly colored, usually banded, usually biconic; spire medium to tail; aperture moderately broad to broad; lip thick- ened, smooth or denticulate, often with stronger or single posterior denticle; external varix present; siphonal notch absent; columella with 4 strong plications occupying more than half the aperture. Type 2 animal; siphon long; man- tle smooth or weakly papillose, extending over external shell surface. Description: Shell (figure 70) small to medium in size (adult length 2.4-12.0 mm). Color translucent to subo- paque, white, cream, to brown, usually with 1-4 narrow, yellow or brown bands. Shell surface smooth, glossy, rare- ly with weak to distinct axial costae. Shape narrowly to broadly biconic, rarely narrowly subovate; weakly to strongly shouldered. Spire height medium to tall, rarely stepped. Aperture moderately broad to broad, wider an- teriorly. Lip thin to moderately thickened, rarely strong- ly thickened, smooth to denticulate, often with stronger or single denticle at posterior fourth of lip, with a distinct external varix. Siphonal notch absent, rarely very weakly indicated; posterior notch usually absent, at most weakly indicated. Ventral callusing usually absent. Columella with 4 strong continuous plications occupying more than half the aperture length. Internal whorls unmodified. External anatomy: 7 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle smooth or weakly papillose, extending over external shell surface, often nearly completely covering shell; foot moderately narrow to moderately broad, about 1-1 '/2 X shell width, and 1-1 '/2 X length; animal variously spotted or blotched with black, opaque white, or red, or with opaque white lines. Penis with sharply pointed distal or posterodistal appendage. Internal anatomy (figure 33): 2 species studied. As in tribe. Proboscis pointed distally; paired salivary glands ascinous or tubular, contained within proboscis, ducts free. Radula: Non-radulate, completely lacking buccal mass, including odontophore. 9 species known or strongly sus- pected to be non-radulate. Distribution and Habitat: Neozelanic (10 species), S. Australian (8 species), Indo-Pacific (28 species), E. Pacific (4 species), W. Atlantic (15 species), W. African (2 spe- cies). South African (3 species). Intertidal to 1,300 m. Fossil Record: Eocene of France, Eocene to Pleistocene of W. Atlantic, Oligocene to Pliocene of Australia, Mio- cene to Pliocene of W. Pacific, to Recent. Nomenclature, Synonymy: Roth (1978;12) synony- mized all of the above genus-group names, a view with which we agree. Powell (1971:223) had previously syn- onymized Longinella under Volvarinella. A cotype of M. maugeana was figured by Kaicher (1981:#2628), who Page 96 THE NAUTILUS, Vol. 109, Nos. 2 & 3 placed it in Dentimargo. treating Longinella as a sub- genus. The type of M. eburneola was figured in Olsson and Harbison (1953:pl. 30, fig. 8). The characters sepa- rating these groups are viewed as specific differences only. The gender of Dentimargo. based on the Latin noun margo can be treated as either masculine or fem- inine. Because the combination "Dentimargo dentifera" used by Cossmann (1899:pi. 4, fig. 15) is generally con- sidered shorthand for Marginella (Dentimargo) denti- fera, there is no direct evidence of intended gender for Dentimargo by Cossmann [(ICZN Art. 30(a)(1)]. Thus, the name is to be treated as masculine. Remarks: This is a widely distributed group, both spa- tially and temporally. Eventually, a thorough revision may enable species groups to be delineated. Until a thor- ough knowledge of the phylogeny is known, useage of informal species groups is a better solution than recog- nition of subgenera. Genus Eratoidea Weinkauff, 1879 (figure 71) Marginella. section Marginella. gruppe Eratoidea Weinkauff, 1879:140 Type species: Marginella margarita Kiener, 1834; SD Cossmann, 1899:87 (figure 71) Diagnosis: Shell minute to medium, white or uniformly colored, rarely banded, broadly biconic; some species axially costate; spire low to medium, often stepped; lip thickened, denticulate; external varix present; siphonal notch absent; columella with 4 strong plications occu- pying half or more of the aperture. Type 2 animal; siphon long; mantle papillose, extending over external shell sur- face. Descriplion: Shell (figure 71) minute to medium size (adult length 1.9-11.0 mm). Color usually translucent to subopaque white, less commonly deep rose, rarely trans- lucent white with 2 narrow brown spiral bands. Shell surface smooth, glossy, some species with weak to distinct axial costae. Shape broadly biconic, weakly to strongly shouldered. Spire low to medium height, often weakly to distinctly stepped. Aperture moderately narrow to broad, not usually distinctly wider anteriorly. Lip weakly to strongly thickened, weakly to distinctly denticulate, with a distinct external varix. Siphonal notch absent; posterior notch usually absent to at most weakly indi- cated. Shell without evident ventral callusing. Cyolumella with 4 strong continuous plications occupying half to more than half the aperture length. Internal whorls un- modified. External anatomy: 2 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle papillose with very long papillae in E. hematita (Kiener, 1834), ex- tending almost completely over external shell surface, undetermined in E. margarita (Kiener, 1834); foot mod- erately narrow to moderately broad, about 1-1 '/2 X shell width, 1-1 '/2 X length; animal variously dotted or spotted with brown, yellow, or opaque white. [Based on pers. obs. of photographs ex. W. R. Liltved and R. Lipe]. Internal anatomy: Unknown. Radula: We have attempted routine extraction of rad- ulae from 2 specimens of E. margarita and found no radula. Based on this and conchological affinities with other non-radulate genera, we strongly suspect that this genus is also non-radulate. Distribution and Habitat: W. Atlantic (9 species). Re- corded from 1 to 1,470 m. Fossil Record: Miocene of Caribbean, to Recent. Nomenclature, Synonymy: Although Weinkauff (1879) used the single genus Marginella. he proposed an outline classification that included the new "gruppe Eratoidea. His "section" apparently corresponds to subgenus, whereas "gruppe" corresponds to section. This genus was placed as a synonym of Marginella (Gibberula) by Coan (1965:189). Gibberula is a cystiscid genus completely unrelated to Marginella or Eratoidea. Remarks: The 4, strong, continuous plications occu- pying more than half the aperture, lack of an anterior notch, and strongly denticulate lip serve to distinguish this group and to place it in Marginellini. The type spe- cies has variably low, weak axial costae, relating it to the smaller axially costate species. t Genus Eurijentome Cossmann, 1899 Cryptospira. subg. Cryptospira. section Euryentome Coss- mann, 1899:95 Type species: Marginella crassilabra Conrad, 1833 (non Marginella crassilabra Bory de St. Vincent, 1827), = Marginella silabra Palmer, 1937 (nom. nov.), = Mar- ginella anatina Lea, 1833 f; OD Diagnosis: Shell medium size; lip strongly thickened, strongly denticulate on its sharp inner edge; strong ex- ternal varix present; weak siphonal notch present or ab- sent; very deep posterior notch present; columella with 4 continuous plications, often with weak 5th plication, and 4 to 5 parietal lirae in some species. Fossil Record: Eocene of Alabama and Mississippi, to Miocene of Trinidad. Nomenclature, Synonymy: The correct name for the type species is M. anatina Lea, 1833, the next available name for the junior primary homonym M. crassilabra Conrad, 1833. The replacement by Palmer (1937:416- 417) was unnecessary and is therefore a junior synonym. Lea's type of M. anatina (fide Palmer & Brann, 1966: 618) was figured in Palmer (1937:pl. 89, fig. 10). Remarks: The presence of parietal lirae initially caused us to consider placement in the (Jystiscidae, but exam- ination of a shell of E. columba (Lea, 1833) revealed G. A. Coovert and H. K. Coovert, 1995 Page 97 unmodified internal whorls with 4 continuous plications. The 4 strong plications, a very strong posterior notch, and probable relationship to S.E. U.S. Eocene Dcnti- margo cause us to place this genus in the Marginellini. t Genus Simplicoglabella Sacco, 1890b MargincHa, subg. Glabella, section Simplicuglabella Sacco, 1890b:21 (313) Type species: Marginella (Glabella) taurinensis Mich- elotti, 1847 t; SD (M) Eames, 1952:119 Diagnosis: Shell medium to large, narrowly to mod- erately broadly biconic; spire medium to tall; aperture moderately broad; lip thickened, smooth, not denticu- late; external varix present; siphonal notch very weak to absent; columella with 4 strong plications occupying more than half the aperture. Fossil Record: Miocene of Italy. Nomenclature, Synonymy: Treated as a subgenus of Marginella in Coan (1965:189). Remarks: This group is an extinct lineage related to Marginella, differing in its narrower shape and generally taller spire. Genus Marginella Lamarck, 1799 (figures 34, 72) Marginella Lamarck, 1799:70 t Marginella. subg Glabella, section Dentietiluglabella Sacco, 1890b:25 (317) [TS: Marginella (Glabella) deshayesi Mich- elotti, 1847 f; M] Cuctimis Deshayes, 1830:34 [in synonymy] Poreellana Gray, 1847142 (non Lamarck, 1801) [TS: Valuta glabella Linne. 17.58, OD (M)] Psemtomarginella Maltzan, 1880:108 [TS: P. adansoni Maltz- an, 1880, = Voluta glabella Linne, 1758; M] Type species: Valuta glabella Linne, 1758; M (figure 72) Diagnosis: Shell small to very large, white to colored, always patterned, biconic to obconic, lacking axial costae; spire low to medium height; aperture broad; lip thick- ened, smooth to denticulate; external varix present; si- phonal notch usually present; columella with 4 strong plications occupying more than half the aperture. Type 2 animal; siphon long; mantle smooth, usually not readily extending over external shell surface. Description: Shell (figure 72) small to very large (adult length 3.5-74.1 mm). Color white, cream, tan, gray, or reddish-orange to brown, always with pattern of white to dark dots or spots, spiral lines or bands, or mottled, or with axial streaks or lines, or various combinations. Shell surface smooth, glossy. Shape biconic to obconic, rarely narrowly biconic, weakly to strongly shouldered. Spire low to medium height, rarely tall. Aperture moderately broad to broad, wider anteriorly. Lip moderately thin to strongly thickened, smooth to weakly or strongly den- ticulate, with a distinct external varix. Siphonal notch usually distinct, absent to weak in some species. Posterior notch absent to weak or distinct. Ventral callusing usually absent, strong parietal callus present in some species. Columella with 4 strong continuous plications occupying more than half the aperture length. Internal whorls un- modified. External anatomy: 24 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle smooth, ap- parently not readily extending over external shell surface in most species; foot moderately broad, about 1 '/2 X shell width, and 1-1 '/2 X length; animal variously spotted, mottled, streaked, or lineated with white, black, red, or yellow on a translucent or variously colored background. Internal anatomy (figure 34): 3 species studied. As in tribe. Proboscis rounded or blunt distally; paired salivary glands tubular, free, ducts attached to walls of esophagus. Radula: Non-radulate, completely lacking buccal mass, including odontophore. 16 species known or strongly sus- pected to be non-radulate. Distribution and Habitat: Indo-Pacific (5 species), W. Atlantic (1 species), W. African (32 species), South Af- rican (25 species). Intertidal to 2,083 m. Fossil Record: Miocene and Pliocene of Italy, to Recent. Nomenclature, Synonymy: The type species of Den- ticuloglabella, M. deshayesi, falls well within the range of Marginella and this genus is here synonymized. For a discussion of the "pseudogenus" Pseudomarginella, see Cooke (1922). Remarks: This group is distinguished by the 4 strong plications occupying more than half the aperture, plus the colored, patterned shell and lack of axial costae. For discussion of the M. musica-diadochus group, see Coo- vert (1989a). Genus Glabella Swainson, 1840 (figure 73) Glabella Swainson, 1840:133, 324. Marginella. section Phaenospira Hinds, 1844:72 [TS: Margi- nella noduta (sic.) Hinds, 1844, = Marginella nodata Hinds, 1844; SD (M) Gray, 1847:142] Marginella, subg Marginella. section Faba Fischer, 1883:602 [TS: Marginella faba (Linne, 1758), = Voluta falx Linne, 1758; M] Type species: Valuta faba Linne, 1758; SD (M) Gray, 1847:142 (figure 73) Diagnosis: Shell medium to large, white to colored, always patterned, biconic, distinct axial costae present; spire medium to tall; aperture narrow to moderately broad; lip thickened, distinctly denticulate; external varix present; siphonal notch present; columella with 4 strong plications occupying more than half the aperture. Type 2 animal; siphon long; mantle extension undetermined. Page 98 THE NAUTILUS, Vol. 109, Nos. 2 & 3 Description: Shell (figure 73) medium to large (adult length 6.2-38.0 mm). Color white, gray, or yellowish- olive to olive in background, always with pattern of spiral lines or rows of dots, a.xial lines or streaks, or variously mottled, spotted, or streaked. Shell surface smooth, glossy, with weak to usually distinct axial costae. Shape broadly to narrowly biconic, weakly to strongly shouldered. Spire medium to tall. Aperture moderately narrow to mod- erately broad, usually widest medially. Lip moderately to strongly thickened, distinctly denticulate, with a dis- tinct external varix. Shell with siphonal notch weak or distinct. Posterior notch absent to weak or distinct. Shell usually without evident ventral callusing. C.olumella with 4 strong continuous plications occupying more than half the aperture length. Internal whorls unmodified. External anatomy: 2 species studied. Type 2 animal; tentacles long, slender; siphon long; mantle extension undetermined; foot broad, about l'/2 X shell width, l'/2 X shell length; animal variously spotted, blotched, or streaked with yellow or reddish-brown. [See summary of G. adanwni (Kiener, 1834) in Coovert (1987g:12) and figure of G. lucani (Jousseaume, 1884) in Gofas & Bran- dao (1985:85, fig. b)]. Internal anatomy: Unknown. Radula: Non-radulate, completely lacking buccal mass, including odontophore. 1 known non-radulate species (Coan & Roth, 1976:220). Distribution and Habitat: Indo-Pacific (4 species), W. African (13 species). Intertidal to 183 m. Fossil Record: Known only from the Recent. Nomenclature, Synonymy: Various authors have des- ignated different type species, including M. prunum (Gmelin, 1791) by Fischer (1883:602) and M. bifasciala Lamarck, 1822 by Redfield (1870:221), but the desig- nation of M. faba by Gray (1847:142) was the earliest. A photograph of the type of G. faha has been studied (pers. obs. ex. photograph of P. W. Clover). The type of G. nodata was figured by Kaicher (1981:#2640). Both fall well within the limits of the genus. Phaenospira had previously been synonymized by Coan (1965:189). Remarks: We consider axial costae to be a species-level character in other genera. In this case, though. Glabella seems to form a natural assemblage, albeit closely related to Marginella. Besides possessing axial costae, this group always has a strongly denticulate lip and an aperture that is broadest medially. DISCUSSION Family Relationships Previous sections of this paper outlined the fundamental differences between the two families of marginelliform gastropods — the Cystiscidae and the Marginellidae. Shared similarities between each of these two families and other neogastropod families (Table 5), in particular. characters presumed to be derived, will be examined next. The Cystiscidae have a number of unusual characters that differentiate them from the Marginellidae. Inter- estingly, these same characters are shared with the Oli- vidae. The partially resorbed "cystiscid internal whorls" are characterized by being extremely thin, modified in shape, and having the columellar plications reduced to a single, axially oriented, sharp edge internally (figures 2, 3). A very similar type of internal whorl, though, is seen in members of the genus Oliva, clearly visible in X-ray photographs in Zeigler and Porreca (1969:8) and confirmed by personal observations of both young and adult shells of O. sayana Ravenel, 1834. Similar internal shell construction is found in Agaronia (pers. obs.). In Olivella, the internal whorls are often nearly completely resorbed rather than modified and thinned. For further discussion, see Olsson (1956:169), Kantor (1991:24, 41- 43, 49), and Marcus and Marcus (1959:107-108). The only other gastropod families reported to have internal whorl resorption are Cypraeidae and Conidae, in which whorl walls are greatly thinned, and Neritidae and El- lobiidae, in which the internal whorls are completely resorbed (Moore, 1960:1121; Fretter & Graham, 1962: 69). Because internal whorl resorption is uncommon in gastropods, we consider this to be a shared, derived char- acter between the Cystiscidae and Olividae. The multiplicate columella of cystiscids consists of 1 or 2 anterior columellar plications that are reduced in- ternally, plus parietal lirae posterior to these plications. In the Olividae, true columellar plications are absent, but a sharp, axially oriented columellar edge is found internally. The parietal area often bears distinct lirae. Although similar to the multiplicate columella of cystis- cids, the olivid columella lacks anterior columellar pli- cations. The radular morphology is of particular interest in differentiating the Cystiscidae from the Marginellidae. The long, narrow radula of at least 80 to 200 or more rachidian plates is characteristic of the cystiscids. Lateral teeth are present only in the genus Plesiocystiscus, in which the radula closely resembles those of certain olivids (Olsson, 1956:figs. 9, 10). Radulae and subradular mem- branes of cystiscids were compared with those from: Oliva sayana Ravenel, 1834, Agaronia griseoalba (von Martens, 1897), and Olivella (Pachyoliva) semistriata (Gray, 1839). In all three olivid species, odontophoral cartilage hoods, very similar to those in cystiscids, were found. Separate odontophoral cartilages, each fitting into one of the flanking odontophoral cartilage hoods, were foimd in O. saijana and A. griseoalba. These cartilages are very similar in general shape to those of Pcrsicula (figures 36, 37). These two olivids also have the posterior end of the radula, which extends beyond the odonto- phore, encased in a typical neogastropod radular sac and beginning as a developmental series of nascent plates. Both species have lateral teeth, but are otherwise similar in radular morphology to the cystLscid genus Persicula (figures 13-16). For further discussion of Oliva radular G. A. Coovert and H. K, Coovert, 1995 Page 99 Table 5. Comparison of major taxoiiomic cliaractcrs of marginelliform groups with related iieogaslropod families. See text for further discussion. Character Volutidae Marginel- loninae Marginelli- nae Cystiscidae Olividae Internal whorls rcsorbed no no no YES YES in most Labial lirae absent absent absent PRESENT in some PRESENT in some Siphonal notch YES in some YES in some YES in some YES in some YES Plications continuous continuous continuous multiplicate multiplicate Head with lateral lappets & dorsomcdial channel PRESENT in some PRESENT absent absent absent Shell covered by mantle or foot mantle in some mantle (in Afrivohita) mantle mantle in some; foot in Canalispira foot Marginellid buccal pouch absent PRESENT PRESENT absent absent Ncogastropod radular sac PRESENT absent absent PRESENT PRESENT Odontophoral cartilage hoods 7 absent absent PRESENT PRESENT Radula with laterals YES in some no no YES in Plesiocystise. YES Central rachidian tricuspid YES in some no no YES in 1 genus YES in some Length of radular ribbon moderately short to long moderately short moderately to very short long long (short in Olivella) Odontophoral cartilages separate (in A lei I hoe) 9 fused at least antenorly separate separate or fused anter. Valve of Leiblcin PRESENT PRESENT PRESENT or absent PRESENT PRESENT Gland of Leiblein Ig. sacculate, or w/terminal bulb Ig. sacculate with terminal bulb short gland, no long duct short gland, or w/terminal bulb Duct from Gland of Leiblein posterior to nerve ring posterior to nerve nng posterior or ANTERIOR posterior to nerve ring posterior to nerve ring Anal gland PRESENT PRESENT PRESENT absent PRESENT or absent Development direct/ veliger direct direct direct most veliger Page 100 THE NAUTILUS, Vol. 109, \os. 2 & 3 morphology, see Marcus and Marcus (1959:124-5). Oli- vella has a shorter, broader radula with laterals and sep- arate odontophoral cartilages. Marcus and Marcus (1959: 121, fig. 31) and Kantor (1991:26, fig. 5C) further discuss the radular morphology of Olivella, including the odon- tophoral cartilages. The rachidian plates are tricuspid in Olivella and Jaspidella (Olsson, 1956:169, figs. 9, 10) and their radulae strongly resemble the triserial radulae of some Plesiocystiscus. Plesiocystiscus, the only cystiscid genus with lateral teeth, is considered to be the most primitive group in the family. Possession of lateral teeth is certainly a primitive feature, as is the tricuspid ra- chidian plate in cystiscids and olivids (Kilburn, 1981: 354). In summary, the entire radular morphology of cer- tain cystiscids and olivids are very similar. Many of these features, such as the formation of the odontophoral car- tilage hoods and separate, flanking odontophoral carti- lages, are considered to be shared, derived characters. The common possession of a typical neogastropod rad- ular sac is a shared, primitive character. Additional shell features shared between cystiscids and olivids include the glossy exterior of the shell. The mantle at least partially covers the exterior of the shell in the cystiscid genera Cystisciis, Granulina, and Pugntis. In other cystiscid genera, such as Plesiocystiscus, Gibberula, and Persicula, the shell is apparently not covered by the mantle. In olivids, the foot at least partially covers the shell and affords protection, and therefore, the glossy shell exterior is apparently due to secretion by the foot, not by the mantle. A similar animal has been described for the cystiscid genus Canalispira. A glossy shell is ap- parently convergent in many groups and does not by itself suggest a strong affinity between cystiscids and olivids. A strong siphonal notch is found in the cystiscid genera Persicula and Gibberula and is very characteristic of olivids, but is also found in many other groups. Lirae on the inner surface of the outer lip are found in many species of Persicula, Gibberula. and Canalispira, and also in the olivid genus Olivella. This character is less common than the other conchological characters but still could be due to convergence. A comprehensive study and review of all these features in olivids is needed and a common link between cystiscids and olivids should be searched for in Cretaceous material. Anatomically, both the Cystiscidae and Olividae have a valve of Leiblein and a short gland of Leiblein (with a long, convoluted duct and a terminal bulb in Amalda) that empties into the esophagus posterior to the nerve ring (Ponder, 1973:fig. 3). Roth groups have a typical neogastropod radular sac and lack the marginellid buccal pouch. Both families have species that possess a single accessory salivary gland. These characters are also found in other neogastropods. An anal gland is absent in Gib- berula (Ponder, 1970:71) and present or absent in the Olividae (Kantor, 1991:tables 1,2). Cystiscids have direct development, whereas olivids generally have pelagic lar- vae (Boss, 1982:1018). In summary, it is suggested that the Cystiscidae and Olividae have a shared, conmion ancestrv based on char- acters summarized in Table 5. They have much more in common with each other than do cystiscids and margi- nellids, which share no derived characters. Our recog- nition of two families of marginelliform gastropods re- flects these conclusions. The erroneous placement of cys- tiscids and marginellids together was apparently based on superficially similar columellar plications, which we have shown to be completely different, and on externally glossy shells, which are a convergently derived adapta- tion produced by the extended mantle. A search for the ancestry of the now restricted family Marginellidae first involves a consideration of the most primitive group within the family. Harasewych and Kan- tor (1991:17) suggested that the Marginelloninae may be the most primitive group in the family. This is based on the unspecialized features of the foregut, including a large valve of Leiblein lacking a glandular bypass, and the large gland of Leiblein, which lacks a terminal bulb and empties into the esophagus posterior to the nerve ring. We fully concur with this view. All other margi- nellids show specializations of the foregut, such as posse- sion of a glandular bypass around the valve of Leiblein, development of a long, convoluted duct from the gland of Leiblein which has a terminal bulb, or loss of the valve of Leiblein with the duct from the gland of Leiblein passing through the nerve ring. The features of the head in Marginellona and Afrivoluta, including possession of lateral lappets and a dorsomedial channel, are features shared with many volutids (Harasewych & Kantor, 1991: 13) and are very likely derived characters. The split head in cystiscids is considered to be fundamentally different and unrelated to the dorsomedially channelled head seen in marginellonines. The two genera of the Marginellon- inae were originally placed in the Volutidae based on their large shells. Another typical volutid feature in Af- rivoluta is the placement of the anterior columellar pli- cation which does not border the anterior edge of the columella as in all other marginellids. The presence of a weak to strong siphonal notch in the Marginelloninae is typical of volutids, but is also found in olivids and some cystiscids. All volutids and marginellids have shells with unmodified internal whorls and strong, continuous col- umellar plications (Moore, 1955:430; Dillon, 1981:14; Abgrall, 1981:9). A valve of Leiblein and an anal gland are present in both families (absent in some Marginel- linae). All marginellids and most volutids have a rela- tively short radular ribbon and have lost the lateral teeth. The broad, comb-like, multiscupid rachidian plate seen in many genera of marginellids is similarly developed in Voluta (Weaver and DuPont, 1970:fig. 1). Apparently all marginellids have a mantle covering the external shell surface, a feature seen in some volutids. For these reasons, it is suggested that the Marginelloninae, and thus the rest of the Marginellidae, are allied to the Volutidae (Table 5). The marginellids differ in the presence of a buccal pouch. Thiele (1929:353-356) recognized only three genera in his treatment of the Marginellidae, namely Persicula, Marginella. and Marginellona. These three genera, with G. A, Coovert and H. K. Coovert. 1995 Page 101 their included subgenera and sections, correspond very closely to the classification of the Cystiscidae and the two subfamilies of the Marginellidae presented here. The single exception is Thiele's inclusion of Canalispira in Marginella. which is not surprising due to its superficial resemblance conchologically to certain marginellids. Coan's (1965) classification recognized these three groups in name, but differed in the assignment of genera. Ponder (1973:331) suggested that the Volutomitridae, which appeared in the Late Cretaceous, possibly arose from a common stem with the Marginellidae. Voluto- mitrids often have an operculum and their radula has a very odd wish-bone shaped rachidian plate, frequently with lateral teeth. These features, along with a variously sculptured shell, are very different from the marginellids. We believe that this group has little in common with the Marginellidae. Part of the difficulty in determining familial relation- ships lies in the abundance of convergent characters within Neogastropoda (Ponder, 1973:302). An overall tendency toward reduction of the number of radular teeth and their cusps is seen throughout the Neogastropoda. Loss of the lateral teeth occurs in some Mitridae, Volutidae, Volutomitridae, Cancellariidae, as well as in Marginel- lidae and Cystiscidae. Complete loss of the radula occurs in at least some Coralliophilidae, Cancellariidae, Turri- dae, Terebridae, Colubrariidae, and Marginellidae. The development of a long, convoluted duct and a terminal bulb in the gland of Leiblein is seen in several families, including some Muricidae, Olividae, Volutidae, Margi- nellidae, and the Conoidea. The reduction or complete loss of the \ aKe of Leiblein is less common, but is seen in some Mitridae and Marginellidae, and all Harpidae and Conoidea. Besides sharing these last two character states, the Conoidea and the Marginellidae also share the development of a buccal pouch and the passage of the duct from the gland of Leiblein through the nerve ring. All of the above characters shared by the Conoidea and Marginellidae are most likely convergent. The parallel evolution of the Conoidean "poison gland" and the gland of Leiblein in the Marginellidae was discussed by Ponder (1970:77-80), Graham (1966:146), and Fretter (1976:333- 334). More research on the phylogeny of these groups is needed, as well as more study into the nature of the gland of Leiblein in the Marginellidae and its possible use as a "poison gland" (Fretter, 1976:333-334). Phylogeny of the Cystiscidae Our hypothesis of the relationships of the higher taxa within the family Cystiscidae is summarized in figure 74. Presumed ancestral character states include a Type 2 animal with a cystiscid Type 1 radula, and a shell with cystiscid internal whorls, a multiplicate columella, and lacking an external varix. Additional characters include lack of a siphonal notch, presence of a valve of Leiblein, and presence of a small gland of Leiblein with a short duct emptying into the esophagus posterior to the nerve ring. All these shell and radular characters are present T>pe J radula Extcmal vanx present Columella with 2 inlcmally reduced plicauons plus 1 or 2 pseudo-conunuous plicaLons Modified cysQscid iniemal whorls CYSTISCIDAE Type 2 anunal Type 1 radula Columella mulupiicale Cysiiscid imemal whorls Noexlemal vanx Figure 74. Intuitively derived phylogeny of the Cystiscidae. Characters at base are primitive for the family; all others are major specializations for each lineage. in the Plesiocystiscinae, here considered to be the most primitive group in this family. The presence of lateral teeth in the radula in this group, undoubtedly a primitive character state, is most significant. Some species of Ple- siocystiscus have a tricuspid rachidian plate, another primitive feature (Kilburn, 1981:354). Because all other c\stiscids have a greater number of cusps along the ra- chidian plate, an increase in the number of cusps is con- sidered to be a specialization in this family. The re- mainder of the family have lost the lateral teeth, a de- rived condition also occurring in several other neogas- tropod families. The unique character states that distinguish the Cys- tiscinae are a Type 3 animal and a Type 2 radula. Extra is thought to be a recent divergence from Crithe. The extinct genus Topaginella, which has a shell with fine axial costae, is likely an early offshoot from this lineage. Persiculinae are distinguished by the presumed de- rived character states of a Type 4 animal, a Type 3 radula, presence of a siphonal notch, and presence of apertural lirae in many species. As Canalispira is thought to be a recent divergence, the absence of a siphonal notch Page 102 THE NAUTILUS, Vol. 109, Nos. 2 & 3 MARGINELLINAE UJ < z z o _l -I UJ z C5 < Modified Type 6 radula, or Type 5 radula Type 8 or 9 I radula I a o E (- CO 3 Valve of Lciblcm y Loss of valve of Lciblcm Dud from gland of Leiblcin passe: Uirough nerve nng Gland of Lciblein wiih lermmal bulb and convoluted duel Type 6 radula. Type 2 animal Type 7 radula. Type 1 animal Gland of Lciblein with shon duct, without terminal bulb Valve of Lciblein present. without bypass lube MARGINELLIDAE Figure 75. Inliiitively derived phylogeny of the Marginellidae Characters at base are primitive for the family, all others are major specializations for each lineage. in this genus is hypothesized to be a secondary loss of this character. If absence of a notch is considered to be a retained primitive character, then development of a notch in the much earlier Gibberula and Persicula would have had to occur twice, which seems less likely. Can- alispira also has a highly modified animal, lacking a siphon and tentacles. Gibberula is distinguished by the development of a glandular tube bypassing the valve of Leiblein, a derived character (figure 79). Development of an external varix in some species of Persicula is also a derived character for this family. The Cranulininae is, in some ways, the greatest de- parture in the family. Specializations in this group in- clude the very distinctive Type 4 radula with modified odontophoral cartilage hoods, presence of an external varix, presence of modified cystiscid internal w horls, with the columella possessing 2 internally reduced plications in addition to 1 or 2 pseudo-continuous plications. The mantle covers the shell in this group, apparently another specialization within this family. Primitive characters that are retained include a Type 2 animal and the ab- sence of a siphonal notch. The Granulininae is thought to have diverged from the main branch of the family (figure 74) after the lateral teeth were lost from the radula. Because this group has a fossil record dating back only to the Miocene, it could also have diverged from the plesiocystiscine line (which shares a Type 2 animal) after the cystiscines and persiculines diverged, but this would imply that lateral teeth were lost twice in this family. Pugniis is considered to be a recent offshoot from Granulina. Although the Granulininae differs in a num- ber of characters from the other subfamilies, the Cystis- cinae and the Persiculinae differ in their radically dif- ferent animal types. Thus, we feel that all four groups should be accorded equal ranking as subfamilies. Phylogeny of the Marginellidae Our hypotheses of relationships of the higher taxa within the family Marginellidae are summarized in figures 75 and 80. Assumed ancestral characters for this family include a Type 1 animal with a marginellid Type 7 radula, a gland of Leiblein with a short duct and lacking a terminal bulb, and presence of a valve of Leiblein (figure 80). Additional characters include a shell with unmodified internal whorls, continuous columellar pli- cations, and presence of an external varix. All these an- cestral characters have been retained in the subfamily Marginelloninae, which is considered to be the most primitive group in the family. The probable relationship between the Marginelloninae and Volutidae has been discussed previously. Because a number of marginellid groups with specializations of the foregut possess fewer cusps along the rachidian plate, loss of cusps, rather than an increase in cusps (as in the Cystiscidae), is considered to be derived within this family. Further evidence of this is found in the greatly reduced radula of Hydrogi- nella. a highly specialized genus, and eventual loss of the radula in two lineages of the Marginellinae. The Marginelloninae is composed of only two genera, Afri- vohita and Marginellona, which can be easily differ- entiated by the number and orientation of columellar plications. Lack of e\es, a de\ elopment for life in deep water, and a very weak siphonal notch further distinguish Marginellona. The remainder of the family is grouped together as the subfamily Marginellinae, based on the shared spe- cializations of the gland of Leiblein, which ends in a terminal bulb and has a long, convoluted duct (figure 80). The animal is the generalized Type 2 with a Type 6 radula. The rachitlian plate of the Type 6 radula has fewer cusps than the Type 7 radula. The extinct genus M yobaruni was prolwbK an early divergence from this main stem of the Marginellinae but could be ancestral to the entire family, based on its appearance in the late Gretaceous. The Marginellinae diverged at this point into two main lineages. The Austroginellini retained the valve of Leiblein but developed a bypass tube (figure 80). Ad- ditional characters of the Austroginellini, which are ap- parentU primitive, include attachment of the paired sal- ivary glands to the esophagus in front of the vaKe of Leiblein, and the ducts from these glands being embed- ded in the esophageal walls (figure 80). The other lineage. G. A. Coovert and H. K. Coovert, 1995 Page 103 X MiKijrii.'d Tviv 6 radula Type 5 radula X / "Serrala "Austroginella Group" Group" V Type 6 radula 5 columellar phcalions AUSTROGINELLINI Figure 76. Intuitively derived pliylogeiiy of the Austroginellini. Characters at base are primitive for the tribe; all others are major specializations for each lineage. composed of the Prunini and Marginellini, lost the valve of Leiblein, but the duct from the gland of Leiblein passes through the nerve ring and empties into the extreme anterior end of the buccal cavity (figure 80). The Prunini and Marginellini are considered to be sister groups based on these shared features, with the Prunini developing an esophageal caecum, a derived character diagnostic for that tribe. Additional characters of the Prunini / Mar- ginellini lineage, which are probably also primitive, in- clude the paired salivary glands being free of the esoph- agus, and their ducts either attached to the esophagus or free, but not embedded in the walls as in the Austrogi- nellini. The Marginellini is characterized by the complete loss of the buccal mass, including radula and odonto- phoral cartilages, plus the loss of the buccal pouch (figure 80). This loss also occurred in Hyalina, an undoubted member of the Prunini based on its possession of an esophageal caecum as well as shared conchological char- acters (figure 80). The loss of a radula is therefore con- sidered to have occurred in two independent lines, not three as speculated by Coan and Roth (1976:220). Intuitively derived phylogenetic relationships within the tribe Austroginellini are summarized in figure 76. Primitive anatomical characters for this tribe have al- ready been mentioned. Additional conditions presumed to be ancestral include a Type 6 radula and a shell with five plications and a denticulate lip. This tribe diverged into two main lineages at this point. The "Serrata Group developed a gradually weakening modified Type 6 rad- ula, while the "Austroginella Group" developed a Type 5 radula, which is a narrower, fewer-cusped, chevron- shaped modification of the original Type 6 radula. (In- formal groups are preferred at this level to avoid further inflation of ta.xonomic groupings). Within the "Serrata Group," the genus Serrata retained the modified Type 6 radula and most species retained the denticulate lip and at least a weakened fifth columellar plication. Con- uginella, an extinct Miocene genus, lost the fifth plication and developed a distinctive conical shape and posteriorly produced lip. Strotnboginella, another extinct genus, di- verged in the Pleistocene from the lineage with anteriorly crowded columellar plications and lost the fifth plication. Hijdroginella, also in this lineage, continued the trend of anterior crowding of plications, with the subsequent reduction to three plications. Hydroginella also contin- ued the trend of radular reduction, resulting in a greatly reduced Type 9 radula. An additional specialization for this group is the presence of a collabral parietal callus Page 104 THE NAUTILUS, Vol. 109, Nos. 2 & 3 my speci Muiile pustulo^ or papillose many specie Type 6 radut3 Esophageal caaum present I Figure 77. Intuitively derived phylogeny of the Prunini. Char- acters at base are primitive for the tribe; all others are major specializations for each lineage. ridge just posterior to the columellar plications, in which the anterior end often superficially appears to be a weak fourth plication. This derived character, along with the distinctive radula, distinguishes this well-defined genus. One species is a known fish parasite. We anticipate that additional species may also share this behavior. Shell shape, as in many groups, is variable and accounts for the many synonyms of Hydroginella. The genus Serra- taginella has a very distinctive and unusual Type 8 rad- ula. The rachidian plate possesses secondary cusps not seen in any other marginellid, giving the main cusps a serrated appearance. Unusual derived shell characters include two collabral callus ridges, one of which is con- fined to the aperture, and replacement of the fifth pli- cation by a weak denticle. The " Austroginella Group," with a Type 5 radula, radiated into several genera. Pos- session of five strong cohmiellar plications occupying more than half the aperture is ancestral within this group. Protoginella retained the five columellar plications and developed a strong posterior notch. An early offshoot of this branch is the extinct Eocene genus Mioginella, which I 01 Bilurcaicd plications usually conncfietl to collabial callus ridge disially \ Labial denliculaliun present and sut>cqual Shell tjToadly biLomc X Siphonal notch ahscnl Terminal bulb o( gland of Lciblcui rounded apicill' \ Siphonal notch usually piesenl Terminal bulb of gland of Leiblein pointed spically MARGINELLINI U)ss of raijul3. buccal mass, and buccal pouch Figure 78. Intuitively derived phylogeny of the Marginellini. Characters at base are primitive for the tribe; all others are major specializations for each lineage. shares the posterior notch but has carinate shoulders and a weak siphonal notch. The remainder of the "Austro- ginella Group" lost the fifth columellar plication. Niidi- faha is an Eocene offshoot from the Alaginella line. Alaginella is characterized by a weakly to strongly alate posterior corner of the aperture, a strongly narrowed shell anteriorly, and general loss of labial denticulation in most species. This group also lacks distinct siphonal and posterior notches. Variable shape and shoulder de- velopment in this genus is responsible for the numerous genus-group synonyms. Hiwia is an extinct Eocene off- shoot with a distinctive shell having strong axial costae and angulate or carinate shoulders. Austroginella is char- acterized by strong posterior and siphonal notches, as well as a distinctively thickened, smooth lip. Mesoginella is variable in shell shape and other features, has a weak or absent posterior notch, and the fourth plication is often remote. From this lineage diverged a group of genera with an immensed spire and thin, sharp, anteriorly crowded columellar plications. Closia and Ovaginella retained four plications. Strong labial denticulation was retained in Closia, which developed a colored shell, whereas Ovaginella generally lost labial denticulation G. A. Coovert and H. K. Coovert, 1995 Page 105 MARGINELLINAE 2 Marginellona MARGINELLONINAE 80 Figure 79. Intuitively derived phylogeny of the Cystiscidae, subfamily Persiculinae as illustrated by features of the f oregut Character states: 1: Valve of Leibiein without bypass tube, 2: Valve of Leiblein with bypass tube, Cibbenila sp, after Ponder (1970), Persicula interruptolineata after figure 35 herein Figure 80. Intuitively derived phylogeny of the Marginellidae as illustrated by features of the f oregut Character states: 1: Gland of Leiblein with short duct emptying posterior to nerve ring, valve of Leiblein present; marginellid buccal pouch present; marginellid radulae present, 2: Gland of Leiblein with terminal bulb and convoluted duct. 3: Valve of Leiblein with bypass tube; paired salivary glands attach to esophagus just anterior to valve of Leiblein; salivary ducts embedded in esophageal walls, 4: Loss of valve of Leiblein; duct from gland of Leiblein passes through nerve ring; paired salivary glands completely free of esophagus; salivary ducts attached or free, but not embedded in esophageal walls, 5: Esophageal caecum developed. 6: Loss of radula, buccal mass, and buccal pouch Marginellona gigas ulier Harasewych and Kantor (1991), Austroginella johmtoni after Ponder and Taylor (1992), Prunum aff. aletes after figure 31 herein, Hijalina pallida after figure 32 herein, Marginella glabella after figure 34 herein. but did not develop shell coloration Balanctta, with a narrower shell, lost the fourth columellar plication. The phylogeny of the tribe Prunini is summarized in figure 77, Anatomical features of this group have already been discussed, the most significant being the develop- ment of an esophageal caecum, a derived character state unique to this tribe. The Type 6 radula is considered primitive and is retained in nearly all genera. The widely distributed Volvarina lineage is characterized by loss of the external vari.\ and development of a pustulose or papillose mantle, both occurring in many, but not all, species. The Prunum lineage, largely a western Atlantic radiation, has several extant Indo- Pacific species and two genera that are offshoots. Cryptospira developed addi- tional columellar plications in some species, and has a variously colored or patterned shell. Rivomarginella, the only freshwater group, is allied to the Prunum lineage based on the ontogenetic development of its varix. Bul- lata is a relatively recent offshoot of Prunum and is confined to the western Atlantic coast of South America. Much more work is required to fully characterize Prun- um and Volvarina. Attention should be focused on anat- omy, including the odontophoral cartilages and radulae, external features of the animal, including coloration and mantle characters, and shell characters. Study of external varix development shows great potential and involves Page 106 THE NAUTILUS, Vol. 109, Nos. 2 & 3 cutting and polishing shell cross-sections. The final branch of the tribe Prunini is the genus Hyalina, characterized by the complete loss of the buccal mass, including odon- tophore, radula, and buccal pouch. This genus is further characterized by a thin shell and columellar plications that are reduced to two or three in some species. Our reconstruction of the phylogeny of the tribe Mar- ginellini is summarized in figure 78. This group is char- acterized by complete loss of the buccal mass and buccal pouch, and lacks the esophageal caecum of the Prunini. Shell features include four strong columellar plications spanning half or more of the aperture. One branch of this tribe is characterized by absence of a siphonal notch, and the gland of Leiblein has the terminal bulb rounded distally. Further anatomical features in Dentimargo in- clude a distally pointed penis and possession of three glandular structures or seminal receptacles that join the pallial oviduct between the albumen gland and capsule gland. Stazzania is an extinct Eocene offshoot of the Deritimargo lineage characterized by the bifurcated ends of the columellar plications joining a collabral callus ridge. Dentimargo shells usually have a prominent posterior labial denticle that is characteristic. The shells of Den- timargo are also generally narrower and differ in pli- cations, which are usually not bifurcated in Recent spe- cies. Where the plications are bifurcated, the ends do not join a collabral callus ridge as they do in Stazzania. Eratoidea is a Caribbean offshoot with subequal labial denticulation and often weak to distinct axial costae. Euryentome is an extinct Eocene group from the western Atlantic with a strong posterior notch and often has pa- rietal lirae posterior to the columellar plications. The second branch of the tribe has the siphonal notch usually present, and the terminal bulb of the gland of Leiblein is pointed distally. Further anatomical differences that may differentiate this group from the Dentimargo line include a simple penis and only two glandular structures between the albumen gland and capsule gland. Further work is needed to confirm this in more species. Simpli- coglabella is a Miocene offshoot with a narrower shell and a taller spire. Marginella is a large genus with a smooth shell, whereas Glabella is a well-defined group having distinct axial costae. EXTRALIMITAL GeNERA The family Cryptochordidae was questionably included as an extinct subfamily of the Marginellidae by Ponder and Waren (1988:306). This family was originally named by Pchelintsev, et. al (1960:235) with the genus Cryp- tochorda Morch, 1858 as type genus, and Bticcinopsis stromhoides Herrmann, 1781 as type species (OD) of the genus. Specimens of Cryptochorda stromboides from the Eocene of France were examined and this species is unrelated to either the Cystiscidae or the Marginellidae. This species has a tiny protoconch, a strong siphonal notch, irregularly-spaced axial costae, a weak siphonal fasciole, and the columella is bordered by a sharp edge with no columellar plications otherwise evident. The glossy shells have some parietal callusing and a sharp posterior sinus. These are all characters of the Harpidae, and we would therefore agree with the assignment of Cryptochorda to that family by Palmer and Brann (1966: 616-617). This group can be removed from further con- sideration in the Cystiscidae or Marginellidae. Bouchet (1989:82-83) placed the monotypic genus Ta- teshia Kosuge, 1986 (type species T. yadai Kosuge, 1986, OD) in the Marginellidae and regarded it as a margi- nellid with a primitive radula. We initially regarded this genus as a marginellid based on shell morphology but assumed an error had occurred in associating the radula, which is very muricid-like, a fact noted by Kosuge (1986: 78). We have been informed by Dr. Sadao Kosuge (pers. comm.) that radulae were obtained from three specimens and thus it appears no error occurred. This genus was originally placed in the Olividae by Kosuge. We can only conclude that this species, which has a radula unlike any member oi the Cystiscidae or Marginellidae, belongs elsewhere. Perhaps further anatomical work will clarify the relationship of this enigmatic genus, which, admit- tedly, has a very marginellid-like shell. Argentovoluta Vazquez and Caldini, 1989 (type spe- cies A. bottai Vazcjuez & Caldini, 1989) was mentioned by Poppe and Gotto (1992:215) as possibly belonging to the Marginellidae. The columellar plications are clearly volutid and, therefore, this genus can be removed from further consideration in the Cystiscidae or Marginellidae. ACKNOWLEDGEMENTS Because this research spanned 13 years, it is impossible to list everyone who assisted in some way, and we apol- ogize and thank anyone not specifically mentioned. We would especially like to thank all the subscribers to Mar- ginella Marginalia from around the world, who are col- lectively the supporters of this research, and to whom we dedicate this paper. We especially thank Jan and Lee Kremer, of Algonquin, Illinois, for their help in many ways, including providing specimens, loan of material, and their continuous support of this research; Dean R. Hewish, of Victoria, Australia, for willingly sharing in- valuable photographs of types and other shells, SEM pho- tographs of radulae from a wide variety of Australian species, and many shell specimens; and Robert Lipe, of Seminole, Florida, for generously sharing many speci- mens, photographs, and preserved material. We thank these students of the marginelliform gastropods for their generosity and apologize for dividing their favorite fam- ily in two! We also thank all the curators and collections managers at the various museums visited or from which material was generously loaned: Dr. M. G. Harasewych and Dr. Harald A. Rehder, National Museum of Natural History; Russell H. Jensen, Delaware Museum of Natural History; Dr. George M. Davis, Dr. Robert Robertson, Dr. Gary Rosenberg, Andria (Jarback, and Elana Benamy, The Academy of Natural Sciences of Philadelphia; Dr. Wil- liam K. Emerson and Walter E. Sage, III, American G. A. Coovert and H. K. Coovert, 1995 Page 107 Museum of Natural History, New York; Dr. Kenneth Boss, Museum of Comparative Zoology, Harvard Uni- versity; Dr. James H. McLean, Los Angeles (bounty Mu- seum of Natural History; Dr. Riidiger Bieler, Field Mu- seum of Natural History, Cfiicago; Dr. John B. Burch, Museum of Zoology, University of Michigan; Dr. John E. Rawlins, Carnegie Museum of Natural History; Kathie Way and Julia Freeman, British Museum (Natural His- tory); Dr. Philippe Bouchet and Dr. Serge Gofas, Mu- seum national d'Histoire naturelle, Paris; Dr. Phillip H. Colman, Australian Museum, Sydney; Dr. Eliezer de C. Rios, Museo Oceanografico, Rio Grande, Brazil; Dr. Rob- ert G. Moolenbeek, Zoologisch Museum, University of Amsterdam; Dr. E. Gittenberger and Dr. Hermann L. Strack, Nationaal Natuurhistorisch Museum, Leiden. This research would not have been possible without the generous support and help of numerous individuals. We particularly thank the following for their help in many ways: Dr. R. Tucker Abbott, T. P. Keukelaar van den Berge, Peter Bright, Phillip W. Clover, Dr. Eugene V. Coan, Dr. David T. Dockery, III, Francisco Fer- nandes, Piet Hessel, Stan Jazwinski, Sally Diana Kaicher, Dr. Sadao Kosuge, Dr. Harry G. Lee, W. R. Liltved, Jose Aguiar Morales, Ivan Perugia, Richard E. Petit, Colin Redfern, Dr. Barry Roth, Peggy Williams, John Wolff, and Jack Worsfold. We would like to e.xpress our sincere appreciation to the Conchologists of America who provided a grant in 1988 that made a research trip to museums in the eastern United States possible. Additional support has been pro- vided from generous donors to the Malacological Re- search Fund of the Dayton Museum of Natural History and their contributions are gratefully acknowledged. The authors would also like to thank the editor, M. G. Harasewych, and two anonymous reviewers for their helpful comments wfiich aided in improving the manu- script. There are many other individuals too numerous to list to whom we are greatly indebted for their generous assistance. Numerous colleagues, collectors, and shell dealers provided literature, specimens, photographs, and other information. University and museum librarians generously provided their time and aid. We gratefully thank and acknowledge all these individuals for their help and contributions, and regret that space does not permit a complete listing. We also express our gratitude in memory of the friends and colleagues who have passed away. LITERATURE CITED For more complete collations and dates of publication, see Coovert (1988e). Al:)bott. R T and S. P, Dance. 1982. Compendium of sea- shells, a color guide to more than 4,200 of the world s marine shells. E. P. Button, Inc., NY. x -I- 411 p. Abgrall, R. 1981. Radiologie & malacologie. Xenophora 5:8-9. Adams, H. and A. Adams. 1853 In: 1853-1858. The genera of Recent Mollusca; arranged according to their organi- zation. John van Voorst, London 3 vols [Ivi-.xl, 1-484; 2:1-661; 3:pls. 1-138]. Barnard, K. H. 1962. A new genus in the family Marginel- lidae. Proceedings of the Malacological Society of London :35(1):14-15. Barnard, K H. 1963. The family position of Afrivoluta prin- glci Tomlin. The Journal of Conehology 25(5): 198- 199 Behrens, D W. 1984. A marginellid dilemma. Shells and Sea Life 16(12):240-242. Blackwelder, R. E. 1964. Phyletic and phenetic versus om- nispective classification. In: Phenetic and phylogenetic classification. Systematics Association Publication 6: 17-28. Boss, K. J. 1982. Mollu,sca. In: S, P. Parker (ed.). Synopsis and classification of living organisms. McGraw-Hill, NY, p.945- 1166. Bouchet, P 1989 A marginellid gastropod parasitizes sleep- ing fishes. Bulletin of Marine Science 45{l):76-84. Brandt, R. A. M. 1968. Description of new non-marine mol- lusks from Asia. Archiv fiir Molluskenkunde 98(516):213- 289, pis. 8-10 Brandt, R. A. M. 1974. The non-marine aquatic Mollusca of Thailand Archiv fiir Molluskenkunde 105(1-4): 1-423. Brocchi, G. 1814. Conchiologia fossile subappenina con os- servazioni geologiche sugli Apennini e sul suolo adiacente. Stamperia Reale, Milano. 2 vols. [2:241-712, pis, 1-16]. Bruggen, A, C. van. 1963. A re-e,xamination of the types of Recent Mollusca of the Port Elizabeth Museum. Annals of the Cape Provincial Museums 3:75-81. Clover, P. W. 1974. A new species of Cypraea from West Africa and three new species of Marginellidae from the Indian Ocean. The Journal of Conehology 28(4):213-216, pi. 8. Coan, E. V. 1965. A proposed reclassification of the family Marginellidae. The Veliger 7(3): 184-194. Coan, E. V and B. Roth. 1966, The West American Margi- nellidae. The Veliger 8(4):276-299, pis. 48-51. Coan, E, V, and B Roth, 1976. Status ot the genus Hyalina Schumacher, 1817 (Mollusca: Gastropoda). Journal of Mol- luscan Studies 42(2):217-222. Coleman, N. 1975. What shell is that? Paul Hamlyn, Sydney. 308 p, Cooke, A H 1922, On the pseudo-genus Pseitdomarginella, v. Maltzan. Proceedings of the Malacological Society of London 15(l):3-5, Coomans, H. E. 1969, On the identity of Cryptospira glauca and the related Cryptospira ventricosa, (Gastropoda: Mar- ginellidae), Basteria 33(5-6):85-92, Coomans, H, E. 1972. The genus Pac/ji/fcaf/iron (Gastropoda). Basteria 36(2-5):89-96. Coomans, H. E 1975. Notes on Marginella spryi. Basteria 39(1 -2): 23-27. Coomans, H E 1976. Historic en systematiek van de Mar- ginellidae (Gastropoda). In Malacologische Opstellen, pp. 7-18, pis, 4-5, Coomans, H, E, and P. W, Clover. 1972. The genus Rivo- marginella (Gastropoda, Marginellidae). Beaufortia 20(263):69-75. Coovert, G. A. 1986a, Tvpe species list. Marginella Marginalia l(l):l-3. Coovert, G A. 1986b. Generic group names, Marginella Mar- ginalia l(2):5-7, Coovert, G. A. 1986c. A review of marginellid egg capsules. Marginella Marginalia 1(4):13-19. Page 108 THE NAUTILUS, Vol. 109, Nos. 2 & 3 Coovert, G. A. 1986d. Notes on the genus Cystiscus and its type species. Marginella Marginalia l(5):20-24 Coovert, G. A. 1986e. Notes on the genus C.rithc and its type species. Marginella Marginalia 1(6)25-30. Coovert, G. A. 1987a. The genus Afrivoluta and a review of the type species Afrivoluta pringlei Tomlin, 1947 Mar- ginella Marginalia 2(1): 1-8. Coovert, G. A. 1987b. The Persicu/a cornea group Marginella Marginalia 2(2/3):10-19. Coovert, G A 1987c. Additonal notes on the genus Crithe. Marginella Marginalia 2(5):28-35. Coovert, G. A. 1987d. Extra Extra: read all about it. Margi- nella Marginalia 2(6):36-38, 40-41. Coovert, G. A. 1987e. Addenda to Afrivoluta paper. Mar- ginella Marginalia 2(6):39. Coovert, G. A. 19871. Proper useage of the generic names Haloginella, Hyalina, and Volvarina with a diagnosis of the genus Haloginella. Marginella Marginalia 3(1): 1-7. Coovert, G. A 1987g. A literature review and summary of marginellid external anatomy. Marginella Marginalia 3(2/ 3):8-25. Coovert, G. A. 1987h. Notes on the genus Granulina and its type species. Marginella Marginalia 3(4):26-32. Coovert, G. A. 1987i. The external anatomy of two species of Caribbean Marginellidae. Marginella Marginalia 3(5): 33-37. Coovert, G A. 1988a. Marginellidae of Florida, Part I: Gran- ulina hadria. Marginella Margnialia 4(1): 1-8 Coovert, G. A. 1988b. Type species of the genera Austrogi- nella and Mesoginella and their synonyms. Marginella Marginalia 4(2/3):9-26. Coovert, G. A. 1988c. Marginellidae of Florida, Part II: Prun- um succinea with a discussion of Prunum and Volvarina. Marginella Marginalia 4(5)35-42, pi. 1. Coovert, G. A. 1988d. Taxonomic characters in the family Marginellidae: conchological characters. Marginella Mar- ginalia 4(6):43-47. Coovert, G. A. 1988e. A bibliography of the Recent Margi- nellidae. Marginella Marginalia 5(I-5):l-43. Coovert, G. A. 1989a. The Marginella musica-diadochiis group. Marginella Marginalia 6(3-5): 16-29, pi. 1. Coovert, G. A 1989b. A literature review and summary of published marginellid radulae. Marginella Marginalia 7(1- 6):l-37. Coovert, G A. and H K (Coovert 1987 Preparation of mar- ginellid radulae. Marginella Marginalia 2(4):20-27. Coovert, G. A. and H. K. Coovert. 1990. A study of margi- nellid radulae. Part I: Type 6 Radula, "Prunum / Vol- varina" Type. Marginella Marginalia 8(1-6), 9(l-6):l-68. Cossmann, A. E. M. 1899. Essais de paleoconchologie com- paree Co.ssmann, Paris. Vol. 3, 201 p., pis. 1-8. Cotton, B. C. 1944. Australian margin shells (Marginellidae). The South Au.stralian Naturalist 22(4):9-16, pi. 1. Cotton, B. C. 1949. Australian Recent and Tertiary Mollusca, family Marginellidae. Records of the South Australian Mu- seum 9(2):197-224, pis. 17-20. Dillon, J. 1981. Inside seashells. Oceans 14(4): 12-17. Djajasasmita, M. and H. E. Coomans. 1980. 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Rediscovery of Marginellona gigas (Martens, 1904). with notes on the anatomy and systematic position of the subfamily Mar- ginelloninae (Gastropoda: Marginellidae). Nemouria 37: 1-19. Hedley, C. I90I. A revision of the types of the marine shells of the Chevert Expedition. Records of the Australian Mu- seum 4(3):I21-130, pis 16-17 Hewish, D R 1990 Marginellidae feeding habits; Dracula meets the chainsaw massacre. Australian Shell News 70: I -3 Hewish, D R .uid K L. Gowlett-Holmes. 1991. Mollusc type specimens in the South Australian Museum. 4. Gastropoda: G. A. Coovert and H. K. Coovert, 1995 Page 109 Marginellidae. Records of the South Austrahan Museum 25(l):57-70. Hinds, R. B. 1844. Descriptions of Marginellae collected dur- ing the voyage of H.M.S. Sulphur, and from the collection of Mr. Cuming. Proceedings of the Zoological Society of London, 1844 pt. 12 [= vol. 14]:72-77. Hyman, L. H 1967 The Invertebrates, vol. VI. Mollusca I. Aplacophora, Polyplacophora, Monoplacophora, Gastrop- oda, The coleomate Bilateria. McGraw-Hill, NV vii + 792 p. Johnson, R. I. 1964 The Recent Mollusca of Augustus .Addison Gould. United States National Museum Bulletin 239: i-v, 1-182, pis. 1-45. Jousseaume, F. P. 1875. Coquilles de la famille des margi- nelles. Monographic. Revue et Magasin de Zoologie, ser. 3, 3:164-271, 429-435, pis. 7-8. [Also issued separately, pp. 1-115, pis. 7-8]. Kaicher, S. D. 1981. Card catalogue of world-wide shells. Publ. by the author Pack # 26— Marginellidae, Part II. Cards 2604-2709 (lacking # 2687). Kaicher, S. D. 1992. Card catalogue of world-wide shells. Publ. by the author. Pack # 60— Marginellidae, Part III. Cards 6110-6215. [two # 6189, lacking # 6193]. Kantor, Y. I 1991. On the morphology and relationships of some oliviform gastropods. Ruthenica, Russian Malacolog- ical Journal l(r-2):17-52. Kilburn, R. N. 1981. Revision of the genus Ancilla Lamarck, 1799 (Mollusca: Olividae: Ancillinae). 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Boletin de Faculdade de Filosofia, Ciencias e Letras, L'niversidade de Sao Paulo, Zoologia 22(232):99-188, pis. 1-11. Marcus, E. and E. Marcus. 1968 On the prosobranch Ancilla diniidiata and Marginclla fralcrculus. Proceedings of the Malacological Society of London 38(l):55-69. Moore, H. B. 1955. X-rays reveal the inner beauty of shells. The National Geographic Magazine 107(3):427-434. Moore, R. C. 1960 Treatise on invertebrate paleontology Part I, Mollusca 1. Geological Society of .America, Inc and University of Kansas, .x.xiii -I- 351 p. Murray, F. V. 1970. The reproduction and life history of Microginella minuiissinia (Tenison-VVoods, 1876) (Gas- tropoda: Marginellidae). Memoirs of the National Museum of Victoria 31:31-35. Neave, S. A. 19.39-1940. Nomenclator zoologicus: a list of the names of genera and subgenera from the tenth edition of Linnaeus, 1758, to the end of 19.35. 4 vols. Zoological Society of London Nieulande, F A D. van. 1981. 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Carpenter from the West Coast (San Diego to British Columbia) Geological Society of America, Memoirs 76:i-viii, 1-376, pis. 1-.35. Palmer, K.V. W. and D. C Brann. 1966. Catalogue of the Paleocene and Eocene Mollusca of the southern and east- ern United States. Bulletins of American Paleontology 48(218):471-1057, pis. 4-5. Pchelintsev, V. F., 1. A. Korobkov, N. S. Volkova, V. A. Vos- tokova, L. V. Mironova, N. K. Ovenchkin and A. I Sukhova. 1960. Mollyuski-Bryukhonogie, Osnovy Pa- leontologii. Gosudarstvennoe nauchno-tekhnicheskoe iz- datel'stov literaturv po geologii i okhrane nedr Mockba 4:1-360. Petuch, E. J. and D. M. Sargent 1986 Atlas of the livmg olive shells of the world. C^oastal Education and Research Foundation, Charlottesville, Va. .\iii -I- 2.53 p., •39 pis. Ponder, W. F. 1970. Some aspects of the morphology of four species of the Neogastropod family Marginellidae with a discussion on the evolution of the toxoglossan poison gland. Journal of the Malacological Society of Australia 2(1):55- 81. Ponder, W. F. 1973. The origin and evolution of the Neo- gastropoda. Malacologia 12(2):295-.338. Ponder, W F. and J D. Taylor 1992. Predatory shell drilling by two species of Austroginella (Gastropoda: Marginelli- dae). Journal of Zoology, London 228:317-328, pis. 1-4. Ponder, W. F. and A. Waren. 1988. .Appendix. Classification of the Caenogastropoda and Heterostropha — a list of the family-group names and higher taxa. Malacological Re- view, Supplement 4:288-328. Poppe, G T and Y. Goto. 1992. Volutes. Mostra Mondiale Malacologia, Cupra Marittima, Italy. 348 p., 107 pis Powell, A. W. B. 1971. New Zealand molluscan systematics, with descriptions of new species. Part 7. Records of the Auckland Institute and Museum 8:209-228, 31 pis. Powell, A. W. B. 1979. New Zealand Mollusca, marine, land and freshwater shells. Collins, Auckland xvi -I- 500 p., pis. 1-82. Redfield, J. H. 1852. 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Emerson Department of Living Invertebrates The American Museum of Natural History New York. NY 10024 Mr. Richard I. Johnson Department of Mollusks Museum ol Comparative Zoology Harvard University Canibridg.' MA 02L38 Dr. Aurele La Rocque Department of CeoIog\' The Ohio State University Columbus, OH 43210 Dr. James H. McLean Department of Malacology Los Angeles County Museum ol Natural History 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department ol Mollusks Museiun of Comparative Zoolog)' Ilai-vard University Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department of Malacolog)' Delaware Museum of Natunil Histoiy RO. Box 3937 Wilmington, DE 19807 Dr. Donald R. Moore Division of Marine Geology and Geophysics Rosenstiel School of Marine and Atmospheric Science University of Miami 4600 Rickenbacker Causeway Miami, FL 33149 Dr. Gustav Paulay Marine Laboratoiy University of Guam Mangilao, Guam 96923 Mr. Richard E. Petit PO. Box 30 North Myrtle Beach, SC 29582 Dr. Edward J. Pe'tuch Department ol Cacology F'lorida .Atlantic University Boca Raton, FL 33431 Dr. David H. Stansbery Museum of Zoology The Ohio State Universits' Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum ol C^ouiparative Zoology Harvard University Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology University of California at Davis Davis, CA 95616 SUBSCRIPTION INFORMATION The subscription rate per volume is US $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' pavmeut and sent to: THE NAUTILUS,' PO. Box 7279, Silver Spring, MD 20907-7279, USA. Chanter of address: Please inform the publisher ol your new address at least 6 weeks in advance. All communications shotild include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is published ([uarterly by Trophon (;oiporation, 8911 Alton Parkway, Silver Spring, MD 20910. Second Class postage paid at Silver Spring, MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 7279 Silver Spring, MD 20907-7279 THE €9 NAUTILUS CONTENTS ^^.>^)^o^^^\:;;^::::tn.«, volume 109, Number 4 ISSN 0028-1344 V^„,^rt5Holo, MA 02543 The Evolution of Cold Tolerance in Corbicula fluminea (Bivalvia: Corbiculidae) Ill First Report of the Genus Proadusta Sacco, 1894 (Gastropoda: Cypraeidae) from the Western Hemisphere, with a Description of a New Species from the Eocene of Washington 113 Brian R. Kreiser Jeffry B. Milton Lindsey T. Groves Richard L. Squires Patrick Anseeuw Yoshihiro Goto Perotrochus metivieri, a New Species of Pleurotomariid from the South China Sea (Gastropoda: Pleurotomariidae) 117 Gecrat J. \ ermeij Morpholog)' and Possible Relationships of Ecphora (Cenozoic Gastropoda: Muricidae) 120 Donn L. Tippett Taxonomic Notes on the Western Atlantic Turridae (Gastropoda: Conoidea) 127 ROBERT TUCKER ABBOTT 1919-1995 Dr. R. Tucker Abbott, former Curator of Mollusks at the National Museum of History, Smithsonian Institution, Curator and liolilcr of the Pilsbry Chair of Malacology at the Academy of Natural Sciences of Philadelphia, Assistant Director of the Delaware Museum of Natural History, and founding Director of the Bailey-Matthews Shell Museum, died on November 3, 1995 on Sanibel Island, Florida after a long illness. Dr. Abbott was known to the international malacological communitv as a prolific author of numerous scholarK and popular works on mollusks, and as an editor of THK NAUTILliS, INDO-PACIFIC MOLLUSCIA, and M0N0(;R.\PHS OF MARINE MOLLUSCA. .\n obituar\ , which will include a listing of his publications and malacological taxa, w ill appear in Milume 1 10 of THF NAl'TILUS. THE NAUTILUS 109(4):111-112, 1995 Page 111 The Evolution of Cold Tolerance in Corhicula fluminea (Bivalvia: Corbiculidae) Brian R. Kreiser Jeffry B. Mitton Department of Environmental, Population and Organismic Biology L niversit) of Colorado Boulder, CO 80309-0334 USA Since Corhicula fluminea (Miiller, 1774) was first re- ported in the United States in 1938 (Burch, 1944), it has spread rapidh across the continent. In his rexievv of the distribution of Corhicula in the L'nited States, Counts (19S6) hsted it as occurring in 33 states. At that time, most of the site references were for locations south of 40° latitude. Several workers hypothesized that Corhicula appeared to have reached the limit of its ability to extend its distribution northward due to a lack of cold tolerance {e.g. Graney ei al.. 1980; McMahon, 1982). Several ob- servations supported this hypothesis. Laboratory exper- iments indicated that the absolute lower temperature limit for Corhicula was 2° C (Mattice & Dye, 1976). Likewise, it had been noted that severe cold caused heavy mortalit\ (e.g. Horning & Keup, 1964; Rodgers et al.. 1979). Though Corhicula fluminea has become established in numerous northern environments, these locations are protected from winter temperatures by industrial ther- mal effluents, usually from power plants. As a result, Corhicula is now found in association with power plants in Iowa tEkblad, 1975), Minnesota (Cummings & Jones, 1978), Michigan (Clarke, 1981; French & Schloesser, 1991) and Connecticut (Morgan et al., 1992). Graney et al. (1980) suggested that these thermally protected popu- lations may serve as stepping stones in further northern expansion. One exception to the association w ith power plants was a population reported in Wisconsin on the St. Croix River (Cummings & Jones, 1978). However, there have been no further reports on this location so the suc- cess of this population is unknown. These observations suggest that low temperatures are a major restriction on the distribution of Corhicula in North America. Published reports and personal communications reveal that Corhicula fluminea is spreading from thermal ref- uges and expanding its range beyond the limits that had been imposed by a lack of cold tolerance. Corhicula is quite common in the Great Lakes in Michigan and is now well established in small lakes within the state (Dr. J. Carlton, pers. comm., 1995). Likewise, Corhicula has spread from a power plant in Haddam, Connecticut up the Connecticut River past Hartford. During severe win- ters, these populations suffer heavy mortality. Thus, ther- mal refuges provided by power plants, deep water and possibK spring water inputs are important for the per- sistence of the clam in the Connecticut River (Dr. D. Morgan, pers. comm., 1995; NUSCO, 1991, 1993, 1994, 1995). Corhicula is also appearing in new locations. Free- man and Perkins (1992) reported Corhicula in lakes in Lancaster Co. and in the Cozad Canal in Dawson Co., Nebraska, Furthermore, Corhicula is currently found in large numbers at river and canal locations along the Platte River from Brady to Cozad, Nebraska (M. Peyton, Central Nebraska Public Power & Irrigation, pers. comm., 1994). Corhicula is now resident at Donner Lake, which is at an elevation of 5933 feet in the Sierra Nevada, and is typically covered by snow and ice in the winter (G. Stockwell, Savannah River Ecology Lab, pers. comm., 1994). In Colorado the first recorded population was reported by Nelson and McNabb (1994) at the Cherry Creek Reservoir in Denver. Recently, we learned of an- other reported location at Highline Lake in western Col- orado (Dr. S.-K. VVu, pers. comm., 1995). We have investigated two of these northern locations. In October 1994, one of us (B.K.) examined an irrigation canal in Lincoln Co., Nebraska (T12N, R26W sec. 30). At that time, a dense population, approximately 120 in- dividuals per square meter, was found in a canal holding only about .3 meters of water. A variety of age classes was present, with individuals < 10-15 mm shell length being more abundant than older individuals (25-30 mm shell length). In June 1995 we sampled Highline Lake, which is approximately 30 km northwest ol Grand Junc- tion, CO at an elevation of 4697 feet. We found only a few shells along the shore. However, a substantial con- centration of individuals (about 30 per square meter) was discovered by diving 10-15 meters from shore near a boat launch. Furthermore, these individuals were quite large (30-40 mm shell length) suggesting that this pop- ulation has been established for some time. Information obtained from conversation with fishermen concurred with these observations. Their remarks placed the clams Page 112 THE NAUTILUS, Vol. 109, No. 4 as having been established in this reservoir for at least 8 years, \oucher specimens Irom this site have been de- posited at the University of Colorado Museum. Despite predictions about Corbicula fluminea's dis- tribution limits in North America, this invader continues to move northward. In some of these northern locations, it appears that a thermal refuge is necessary. For ex- ample, along the Connecticut River winter mortality can approach 99%. After a severe winter, populations are primarily composed of juveniles, presumably the off- spring of inhabitants of thermal refuges (NUSCO, 1994, 1995). However, Corbicula is present in other locations without the benefit of an apparent thermal refuge (e.g. the Nebraska irrigation canal & Highline Lake). Unfor- tunateK , we have no data on winter temperatures or any information on the amount of winter mortality experi- enced. Without this information we are only able to speculate on the apparent cold tolerance of these pop- ulations. However, these observations do suggest that this in\ ading species is expanding its range by evolving great- er tolerance to cold. LITERATURE CITED Burch, J. Q. 1944. Checklist of west American mollusks. Min- utes, Conchological Club Southern California 38:18. Clarke, A. H. 1981. Corbicula fluminea, in Lake Erie. The Nautilus 95:83-84. Counts, C L., III. 1986. The zoogeographs and history of the invasion of the United States by Corbicula fluminea (Bivalvia: Corbiculidae), In: Prezant, R.S. (ed.), Proceed- ings of the Second International Corbicula Symposium, American Malacologicai Bulletin, Special Edition No. 2:7- 39. Cummings, S. E. and J A. Jones. 1978 Occurrence of Cor- bicula manilensis Phillipi (sic) in the lower Minnesota River. Journal of the Minnesota Academv of Science 34: 13-14. Eckblad, J. W, 1975. The Asian clam Corbicula in the Upper Mississippi River. The Nautilus 89:4. Freeman, P. W. and K. Perkins. 1992. Survey of mollusks of the Platte River. University of Nebraska State Museum, Final Report. 37 pp. French, J. R. P., Ill and D. W. Schloesser. 1991. Growth and overwinter survival of the Asiatic clam, Corbicula flu- minea, in the St Clair River. Michigan Hvdrobiologia 219:165-170. Graney, R. L., D. S. Cherry, J. H. Rodgers, Jr. and J Cairns, Jr. 1980. The influence of thermal discharges and sub- strate composition on the population structure and distri- bution of the Asiatic clam, Corbicula fluminea. in the New River, Virginia. The Nautilus 94:130-135 Horning, VV. B. and L Keup. 1964. Decline of the Asiatic clam in the Ohio River. The Nautilus 78:29-30, Mattice, J. S and L. L. Dye. 1976. Thermal tolerance of the adult Asiatic clam. In: Esch, G. W. and R W McFarlane (eds.) Thermal Ecology 2:130-135. McMahon, R. F. 1982. The occurrence and spread of the introduced Asiatic freshwater clam, Corbicula fluminea (Muller). in North America: 1924-1982. The Nautilus 96: 134-141. Morgan, D. E., M. Kesser, J. F. Foertch and J. M. Vozarik. 1992. The Asiatic clam, Corbicula fluminea. extends its North American distribution to the Connecticut River (ab- stract only). Page 523. In: Blogoslowski, \V. J. (ed.). Aqua- culture and Environmental Stewardship: Milford Shellfish Biologv Seminar - 1991. Eiuironmental Management 16: 521-529, Nelson, S. M. and C McNabb. 1994. New record of .Asiatic clam in Colorado. Journal of Freshwater Ecology 9:79. NUSCO (Northeast Utilities Service Company). 1991. Status report on Corbicula studies at Haddam Neck Plant. As submitted to the Connecticut Department of Environ- mental Protection on August 15, 1991. NL^SCO Letter No. D04855. 11 pp. NUSCO. 1993, 1992 Annual report on Corbicula studies at Connecticut Yankee. As submitted to the Connecticut De- partment ot Environmental Protection on Januar\ 21, 1993. NUSCO Letter No. D06145. 21 pp. NUSCO. 1994. 1993 Annual report on Corbicula studies at Connecticut Yankee. As submitted to the Connecticut De- partment of Environmental Protection on Januarv 27, 1994. NUSCO Letter No, D07282 19 pp NUSCO. 1995. 1994 .Annual report on Corbicula studies at Connecticut Yankee, .■Ks submitted to the t^onnecticut De- partment of Environmental Protection on Januarv 21, 1995. NUSCO Letter No. D08388. 18 pp. Rodgers, J. H., Jr., D. S. Cherry, K. L. Dickson and J. Cairns, Jr. 1979. Invasion, population dynamics and elemental accumulation of Corbicula fluminea in the New River at Glen Lyn, \'irginia. In: Britton, J. C (ed.), Proceedings, First International Corbicula Symposium, p. 99-110, Tex- as Christian University Research Foundation, Fort Worth, Texas. THE XALTILUS 109(4):113-116, 1995 Page 113 First Report of the Genus Proadiista Sacco, 1894 (Gastropoda: Cypraeidae) from the Western Hemisphere, with a Description of a New Species from the Eocene of Washington Lindsey T. Groves Malacology Settioii Natural Histor> Museum of Los Angeles CA)Uiit\ 900 Exposition Boulevard Los Angeles, California 90U07 V^A Richard L. Squires DepartuuMit ol (leologieal Sciences California State L'nivcrsits 18111 Norcllioff Street Northridge. California 9133()-82f56 USA ABSTRACT A new species of cypraeid gastropod, Proadimta gucdcrtunim n. sp., is reported from the middle lower Eocene ("C;apa\ Stage ) upper part of the Oesceiit Eormation, Thurston C^oun- t\, Washington. This new species was found at two localities where shallow-water marine ileposits are iiiterliedded with rock\ shoreline-forming basalt flows, Froddusta Sacco, 1S94 was pre- viousK known onl\ from the lower Eocene to lower Miocene of Europe, Myanmar (= Burma), and Indonesia. Key wards: Proadiista, C\praeidae, Western Hemisphere, Eo- cene, Washington. INTRODUCTION Only five species of the family Cypraeidae are known from the Eocene of the eastern Pacific region (Groves, 1993; 1994a), and Nucleolaria cowlitziana Groves, 1994a is previously the only true cypraeid known from the Eocene of Washington. The first appearance of the genus Proadiista Sacco, 1894 in the Western Hemisphere is recorded here with the description of Proadusta goe- dertorum Groves and Squires, n. sp. from middle lower Eocene ("C]apay Stage ) strata in the upper part of the Crescent Formation, Thurston County, Washington. Proadiista gocdcrtorum is also one of onl> two species of the genus found anywhere in the lower Eocene, the other being P. chevallieri (Cossmann, 1896) from Ypre- sian deposits of Liancourt, Oise Department, France. STRATIGRAPHY The new species is from the upper part of the Crescent Formation of Arnold (1906) at C;SUN localities 1563 and 1564 (= LACMIP localities 16655 and 16848) at Larch Mountain, just west of Olympia, in the Black Hills area in the Washington Coast Ranges (Figure 1). These lo- calities have been the subject of recent studies by Squires and Goedert (1994; in press) involving molluscan pale- ontology, depositional environment, and geologic age. Fossil-bearing rocks at both localities consist of a thin section of richU fossiliferous and conglomeratic silty mudstone interbedded with basalt. Extrusion of the ba- salt caused shoaling and the establishment of a rock\ shoreline community where gastropod and bivalved mol- lusks lived with colonial corals and abundant coralline algae. Shells were transported a short distance seaward where they were deposited as a matri.x of coquina that infilled spaces between basalt boulders. Many of the shells in the cocjuina are small to minute, and their size pre- vented them from being destroyed during transport. Within the coquina are a few larger shells, like those of the new species, that apparently lived in the shallow- subtidal environment where coquina accumulation took place (Squires & Goedert, 1994; in press). GEOLOGIC AGE Based on mollusks and benthic foraminifera, Squires and Goedert (1994; in press) assigned the rocks at both CSL'N localities 1563 and 1564 to the middle lower Eocene ("Capay Stage"). Clark and Yokes (1936) were the first to recognize this west coast mega-invertebrate stage. The stage name is informal; therefore, it is placed in quotation marks. Givens (1974) modified the use of the "Capay Stage, and it is in this modified sense that the name is used herein. As discussed in Squires et al. (1992), this stage is equivalent to the west coast benthic forminiferal Penutian Stage, as used in the emended sense of Almgren et al. (1988) The "Capay Stage is also equivalent to the middle part of the European Ypresian Stage (Squires, 1987). ABBREVIATIONS Abbreviations used for catalog numbers and/or locality mmibers are: CSUN, C>alifornia State L'niversity, North- ridge; LAC;M, Natural History Museum of Los Angeles County, Malacology Section; LAC^MIP, Natural History Museum of Los Angeles County, Invertebrate Paleon- Page 114 THE NAUTILUS, Vol. 109, No. 4 Figure 1. Inde.x map showing localities of the new species described herein (Type locality = CSUN 1563 [= LACMIP 166.55]). Localities are described in the "Localities Cited" sec- tion. tology Section. Measurement parameters are defined as follows: length = greatest distance between anterior and posterior ends; width = greatest distance between lateral margins; and height = greatest distance between base and dorsum. The classification herein follows that of Schilder and Schilder (1971). SYSTEMATIC PALEONTOLOGY Superfamily Cypraeacea Rafine.sque, 1815 Family (]ypraeidae Rafinesque, 1815 Subfamily Erosariinae Schilder, 1924 Tribe Pustuiariini Schilder, 1932 Genus Proadusta Sacco, 1894 Type Species: Cijpraca (Proadusta) denticulina Sacco, 1894 [= C. (P.) splendens (Grateioup, 1845) var. den- ticulina Sacco, 1894 (not of Grateioup)] by subsequent designation of C^ossniann (1903:156). Lower Oligocene (Lattorhan Stage), Garcare, Liguria Province, north- western Italy [Note: The Lattorhan Stage of Mayer-Eym- ar (1893) has been rejected as a standard chronostrati- graphic unit of the lowermost Oligocene and may span the interval from the lale Middle Eocene to the earliest Oligocene [= Priabonian through earliest Hupelian] (Berggren et al., 1985; Prothero, 1994)]. Original Description: Testa affinis .Adusta, sed spira non excavate, plus minusve protninens (Sacco, 1894:33), Diagnosis: Shell small to medium in size (19 mm), ovate to pyriform, with produced extremities; aperture curved posteriorly, with narrow deep canals; teeth numerous and fine; fossula wide, shallow, smooth (modified from Wenz, 1941). Remarks: Schilder and Schilder (1971) recognized at least 18 species and 10 subspecies of Proadusta from lower Eocene (Ypresian) through lower Miocene (Aqui- tanian) strata of Europe and southeast Asia. (Specific ranges include: EOC^ENE of Belgium, England, France, Germany, Indonesia (Borneo), and Italy; OLIGOCENE of France, Germany, Italy, and Myanmar (= Burma); and MIOCENE of France). Proadusta goedertorum, sp. nov. is the first report of this extinct genus in the Western Hemisphere. Dolin (1991) described Cypraeacites blowi from the upper Eocene (lowermost Jacksonian), Moodys Branch Formation, Montgomery Landing (= Creole Bluff), Grant Parish, Louisiana [U.S. Geological Survey locality 2003]. Groves (1994b) assigned C. bloivi to the genus Proadusta following Schilder (1939) and Schilder and Schilder (1971) who had placed Cypraeacites in synonymy with Proad- usta. Under Article 20 of the ICZN (1985) the genus Cypraeacites is not an available name and blowi is here assigned to the new world genus Cypraeorbis of Conrad (1865) based on similarities of shell shape, terminal ridg- es, fossula, and dentition, therefore establishing P. goe- dertorum as the first record of the genus Proadusta in the Western Hemisphere. The new species of Proadusta described herein from the middle early Eocene of Thurston County, Washing- ton provides additional evidence that tropical conditions existed in this region. Durham (1950) reported that the tropics extended northward of 49°N along the Pacific coast of North America during most of the Eocene. Squires and Groves (1993) also documented an Eocene tropical climate in King County, Washington by the presence of the ovulid species Sulcocypraca mathcwsonii (Gabb, 1869) from the Tukwila Formation. Groves (1994a) fur- ther documented tropical conditions in the Washington Eocene with the description of Muclcolaria cowlitziana from the Cow litz P'ormation. Proadusta goedertorum Groves and Squires, new species (Figures 2-5) Diagnosis: A Proadusta ot small to medium size with produced extremities, posteriori) curved aperture, fine teeth, and smooth, wide fossula. Description: Shell ovate shaped, small to medium in size with prominent produced extremities; spire covered; dor- sum highly arched and smooth; maximum height slightly posterior ol mitl point; slight marginal callus; base convex; aperture curved posteriorly toward columella; labial lip with 26 strong teeth with smooth interstices, coluniellar lip with 17 weak teeth with smooth interstices; teeth L. T. Groves and R. L. Squires, 1995 Page 115 sS Figures 2-5. Proadusta goedcrtorurn Groves & Squires, n. sp., holotype 2. Abapertural view. 3. Apertural view. 4. Left-lateral \ie\v, 5. Right-lateral view [all x 2.57], produced at both extremities torming slight anterior marginal ridges, indistinct in posterior canal; fossula wide, smooth, concave; posterior and anterior terminal canals shallow and narrow; columellar lip slightly inflated; shal- low pit abo\e posterior terminus at base of dorsum. Type Material: Holotype LACMIP 12375, paratype LACMIP 12376. The holotype measures 19.4 mm in length, 12.3 mm in width, and 10.8 mm in height. The paratype measures 17.1 mm in length, 10,4 mm in width, and 8.9 mm in height. Both specimens display original shell material. A third poorlv preserved specimen from CSUN locality 1564 (= LACMIP locality 16848) also displays original shell material Type Locality: CSUN locality 1563 (= LACMIP localit% 16655), Larch Mountain area. Black Hills, Thurston County, Washington (47°59'09N", 123°8T2"W). Middle early Eocene age ("Capay Stage '), upper part of the Crescent Forination. Comparison: The new species is most similar to Proad- usta moloni (Ba>an, 1870) from the middle Eocene (Lutetian Stage) of San Giovanni Ilarione, Veneto Dis- trict, Italy. Proadusta goedertorum has more numerous, finer apertural teeth and less pronounced terminal ex- tensions. The new species also resembles P acyensis (Raincourt, 1876) from upper Eocene (Auversian Stage) of Acy, Oise Department, France but has a less sinuous aperture, narrower posterior and anterior canals, and a wider labial margin. Discussion: The excellent preservation of the new species allovNS for its unequivocal generic assignment. Proadusta goedertorum is significantly different from all other east- ern Pacific cypraeids and is the only representative of this genus in the Western Hemisphere. Etymology: This new species is named after James L. and Gail H. Goedert, Gig Harbor, Washington, for their numerous valuable contributions to the study of inver- tebrate and vertebrate paleontology of Washington. ACKNOWLEDGMENTS Special thanks to James L. and Gail H. Goedert, Gig Harbor, Washington for collecting the type material and for generously donating the specimens to LACMIP. Without their efforts and cooperation, this paper would not be possible. We especially thank Mark Herbert and Don McNamee (LACM Research Library ) for processing numerous inter-library loans. Henry W. Chaney (Santa Barbara Museum of Natural History) arranged for access to the rare-book collection at SBMNH. Suzanne Hen- derson, Melinda Hayes, and Jean Crampon (Allan Han- cock F'oundation, University of Southern California) as- sisted in locating rare and obscure references. James H. McLean (LACM, Malacology). LouElla R. Saul (LAC- MIP), and three anonymous reviewers critically read the manuscript LITERATURE CITED Almgren, A. A., M. V. Filewicz and H. L Heitman. 1988. Lower Tertiary foraminiteral and calcareous nannofossil zouation of C^alifornia: An o\er\ lew and recommendation. In: Filewicz, M. \'. and R. L Stjuires ieds. ), Paleogene Stratigraph\ , West Coast of North America. Pacific Sec- tion, Society of Economic Paleontologists and Mineralo- gists 58:83-105, figs. 1-7, .KriKild, R, 1906, Geological reconnaissance of the coast of the Ol> mpic Peninsula, Washington. Bulletin of the Geo- logical Society of America 17:451-468, pis. 55-58, Bavan, F 1870 Sur les terrains Tertiaires de la X'enetie. Bulletin de la Societc Geologiciue de France (2)27(4):444- 4S7, Berggren, W. A., D. V. Kent, and J. J. FKnn. 1985. Jurassic to Paleogene: Part 2 Paleogene geochronology and chron- ostratigraphy. In Snelling, N.J. (ed.). The chronology of the geological record The Geological Societ\ Memoir 10: 141-195. figs. 1-6 Clark, B. L. and H. E. \okes. 1936. Summary ol marine Eocene sequence of western North .Vmerica Bulletin of the Geological Sociel) ol .America 47:851-878, pis. 1-2. Page 116 THE NAUTILUS, Vol. 109, No. 4 Conrad, T. A. 1S63. Catalnmic ot the Eocene and Oligocene Testatea of the I iiited States. .American Journal of Con- chology Ul,):l-.35, ('ossinann, A. E. M 1.S96. .Vppendice no. 2 an catalogue illustre de.s coquilles fossiles de 1 Eocene des environs de Pari.s.Annaies de la Societe Royale Malacologique de Bel- gique 31:3-94, pis. 1-3. Cossmann, A. E. M. 1903. Essais de Paleoconchologie Coni- paree. ("inquienic h\raisoii Paris. 215 p., 16 figs , 9 pis. Dolin, C. and L. Doiin 1983. Revision des Triviacea et- Cypraeacea (Mollusca, Prosobranchiata) Eocenes recoltes dansies locaiites de Gan (Tuilerie et Acot) et Bosdarros (Pyrenees .\tlantiques, France), Mededelingen van de- W'erkgroep voor Tertiaire en Kwartaire Geologie 20(1): 5-48, figs. 1-31. Doiin. L. 1991. Cypracacites hlowi n.sp., first occurrence of tlie genus in the new world Studies on Paieogene Cy- praeoidea (Mollusca: Gastropoda) from the Gulf Coast Ba- sin-ll. Mississippi Geology ll(4):6-7, fig. 1. Durham,] W 19.50 Cenozoic marine climates of tiie Pacific coast. Bulletin of the Geological Societv of America 61: 1243-1264. figs. 1-3, Gabb, W. M. 1866-1869. Cretaceous and Tertiary fossils. Pa- leontology of California, State Geological Survey 2:1-38 [1866]; 39-299, pis. 1-36 [1869]. Givens, C. R 1974. Eocene inolluscan biostratigraphy of the Pine Mountain area, Ventura C^ounty, C^alifornia. Llniver- sit\ of California Publications in (Jeological Sciences 109: l-i07, figs. 1-7, pis. 1-11, Grateloup, J. P. S. de. 1827. Description de plusieurs de co- quilles fossiles des environs de Da.x (Landes), Bulletin d Histoire Naturelle de la Societe Linneenne de Bordeaux 2:3-25. Groves, L. T. 1993 P'ossil and Recent species of eastern Pacific Cypraeacea (Pediculariidae, Cypraeidae, and Ovulidae). Western Societv of Malacologists. .'\nnual Report 25:11- 14 Groves, L. T. 1994a. New species of Cypraeidae {Mollusca: Gastropoda) from the Miocene of California and the Eo- cene of Washington. The Veliger 37(3):244-252, figs. 1-13. Groves, L. T. 1994b. Catalog of fossil and Recent Cypraeidae and Eocypraeinae (Ovulidae) described since 1971. The Cowry n.s. 1(1):5-16. Mayer-Eymar, K. 1893. Lt- l.iguricn et le Tongrien en fig\ pte (1), Bulletin de la Societe (Jeologicjue de France. 3rtl ser., 21:7-43 Prothero, D. R 1994 The Eocene-Oligocene transition. Par- adise lost. (Columbia L^niversity Press, New York, xvii + 291 p., numerous figs. Rafincsque, C. S. 1815. Analyse de la nature, ou tableau de I'universet des corps organises. Palermo. 224 p. (reprinted 1864; 1984] Raincourt, M de. 1876 Description d especes nouvelles du Bassin de Paris. I5ulletin ile la Societe Getjlogiciue de ('ranee (3)4:290-293. pi. 5, Sacco, F. 1894 1 molluschi dei Terreni Terziarii del Piemonle e della Liguria. Parle W (C^ypraeidae, ed .'\mphiperasi- dae). Carlo (;lau.sen, Torino, Italy. 74 p., 3 pis. Schilder, F. A. 1924. Systemalischer Index der rezenten- CA-praeidae. Archiv fijr Naturgeschichle 90.\(4): 179-214. Schilder, F. A. 1927. Revision der tispraeacea (Moll., (Jastr, ) Archiv fiir Naturgeschichle 91 A( 10)1-171 Schilder, F. A. 1932. Cypraeacea. In W yuenstedt (ed.),Fossilium Calalogus. 1: .Aiunialia, |)t, .55, W, Junk: Berlin: Pp 1-276 .Schilder, FA. 1939. Die genera der Cypraeacea. Archiv fiir Molluskenkunde 71:16.5-201, pis, 7-8. Schilder, M, and F, A Schilder 1971 A catalogue ol living and fossil cowries Instilul Royal ties Sciences .\aturelles de Belgique Menioire 85: 1-246. Squires, R L, 1987 Eocene molluscan paleontology of the Whitaker Peak area, Los Angeles and N'entura C>)uiities, California. Natural History Museum of Los Angeles Coun- ty, Contributions in Science 388:1-93, figs. 1-135. Squires, R. L. and L. T. Groves. 1993. First report of the ovulid gastropod Sulcocypraca mathewsonii (Gabb, 1869) from the Eocene of Washington and Oregon and an ad- ditional report from C^alilornia, The X'eliger .36( 1 ):81-87, figs. 1-4. Squires, R, L and J, L, Cloedert 1994 .New species of earlv Eocene small to minute mollusks Irom the Oescent For- mation, Black Hills, southwestern Washington The X'e- liger 37(3):253-266, figs. 1-29, Sciuires, R. L, and J, L. Goeilert In Press. New species of small to minute gastropods of early Eocene age from the Crescent Formation, Black Hills, southwest Washington, The Veliger, Squires, R. L., J. L. Goedert and K. L Kaler 1992 Paleon- tology and stratigraphy of Eocene rocks at Pulali Point, Jefferson County, eastern Olympic Peninsula, Washington. Washington Division of Geology and Earth Resources, Re- port of Investigations 31:1-27, figs. 1-7, pis. 1-3. Wenz, W, 1941, Superfamily Cypraeacea. /n: Schindewolf. O H (ed) Handbnch der Palaozoologie. Band 6, Proso- branchia, Teil 5. Gebriiiler Bt)rntraeger, Berlin, Germany, p, 949-1014. figs. 2765-2910, LOCALITIES CITED CSUN 1.563 (= LACMIP 16655). At elevation of 680 m (2230 ft. ), exposed in roadcut on northeast side of logging road, 47°59'03"N, I23''8'12"VV, .300 m north and 50 m east of southwest corner of section I, T17N, R4V\', and 500 m S32°E of Larch Mountain, Capitol Peak U.S. Geo- logical Survey 7.5', provisional edition 1986 quadrangle, Thurston County, Washington, 1:24,000. Middle early Eocene ("Capav Stage"), Crescent Formation. Collec- tors: J.L. & G.H. Goedert, 1992-1995. C:SUN 1.564 (= LACMIP 16848). At elevation of .5.30 m, roadcut e.xposure, 800 m (2624 ft.) north and 50 m (164 ft.) west of southwest corner of section 25, T18N, R4W, Summit Lake L'.S. Geological Siirvev, 7.5', 1981 quad- rangle, Thurston C>oimty, Washington, 1:24,000. Middle earlv Eocene ("Capav Stage ), Crescent Formation. Col- lectors: J.L. & G.H. Goedert, 1992-1995. THE NAUTILUS 109(4): 117-1 19, 1995 Pane in Perotrochus metivieri, a New Species of Pleurotomariid from the South China Sea (Gastropoda: Pleurotomariidae) Patrick Anseeuw Mispelstraat 18 9820 Merelbeke BELGIUM Yoshihiro Goto Yamamoto-Cho Miiiami 5-1-6 Yao Citv, Osaka Prefecture 581 japan' ABSTRACT Perotrochus nictivieri. a new species of pleurotomariid, is de- scribed from bathyal depths in the South C^hina Sea It belongs to the thin-shelled group within Perotrochus termed "Group B" b\ Baser (1965) and "Perotrochus africanus - complex" b\' Wagner and Coomans (1990). Comparisons are made with the other species of Perotrochus sensu lata that inhabit this area. Key words: Pleurotomariidae, Perotrochus. new species. South China Sea INTRODUCTION A medium-sized specimen of Perotrochus, live-collected but lacking soft parts or operculum, was first brought to our attention b\ Mr. G. Campanini of Pavia, ItaK , w ho had obtained it indirect!) from Russian trawlers oper- ating in the South China Sea off Vietnam. Identification proved problematic as this specimen resembled to a cer- tain e,\tent both juvenile examples of the high-spired form of Perotrochus africanus Tomlin, 18-18 and large adult specimens of P. vicdani Kosuge 1980. A second specimen was obtained by Mr. Donald Dan of Baltimore, L'SA. The data accompanying this specimen was vague. The shell came from a Russian source and was alleged to have been taken off New Zealand. A third specimen has since been obtained b\' the junior author. Close examination and direct comparison ot these three specimens re\ealed consistent tlifferences in a series of characters in shell morphologs that allow these speci- mens to be distinguished from other Recent Pleuroto- mariidae. This species is herein described as new, even though the localit\ data supplied w ith all three specimens is imprecise. SYSTEMATICS Superfamily Pleurotomarioidea Swainson, 1840 FamiK Pleurotomariidae Swainson, 1840 Genus Perotrochus P. Fischer, 1885 Perotrochus metivieri Anseeuw and Goto, new species (Figure 1) Description: Shell (fig. 1) medium-sized for genus (maximum diameter to 68.0 mm, minimum diameter to 59.4 mm, height to 60.4 mm), thin, light, non-umbilicate. Shell profile trochiform with clearh' gradate spire. Suture adpressed. Mean spire angle 82° (n = 3), profile of pe- riphery rounded, base moderateK- convex. Protoconch damaged or worn on all specimens, glassy, of slightly more than 1 whorl, with weakly flared lip. Teleoconch of up to 7-1- whorls. Selenizone appears near suture in first 1/4 whorl, descends to mid-whorl b\ whorl 2, and below mid-whorl by whorl 5. Selenizone flush with whorl surface or slightK- concave, margins flush or slightly keeled. Anal slit broad ( to 4.4 mm), long, with upper margin spanning up to 100°, lower margin to 72° Axial grow th striae dominant and strongK prosocline between suture and selenizone, weak and prosocyrt between se- lenizone and periphery, opisthocyrt on selenizone. Spiral sculpture of thin cords (10-12 between suture and se- lenizone, 1 on selenizone, 5-6 between selenizone and periphery) well-defined on earlv whorls, creating dis- tinctly cancellate, finely beaded sculpture above and be- low selenizone of first 3-4 whorls. After 4th whorl, spiral cords rapidly become obsolete, persisting only as irreg- idarK-spaced threads in area near the suture and be- tween selenizone and periphery. Aperture subquadrate to o\al, curse of columella gently sigmoidal, columellar margin nacreous, not thickened, extending 1/5 the dis- tance from axis to periphers. L'mbilicus excavated but imperforate. Base weakly convex, with 40-44 fine spiral cords intersected b\' regular, fine, axial growth lines, producing finely cancellate surface. Protoconch and first teleoconch w liorl porcellaneous white; base color of te- leoconch buft w ith occasional darker patches and slight, pinkish suffusion at margins of selenizone. Entire outer surface slightly iridescent due to thinness and translu- cency of outer shell la)er. Interior of aperture (except base of previous whorl) cosered b\' thin nacreous layer, within which the pattern of exterior sculpture is dis- cernible. Nacre covers inner surfaces above and below slit, with extremely thin chamfer of constant width at slit margins. Operculum and soft parts unknown Type locality: Traw led in 350 m in the South China Sea off N'ietnam. Page 118 THE NAUTILUS, Vol. 109. No. 4 Figure 1. Pcwtwchm metwwn Ansefuw and Colo, new species. Apertural, lateral, apieal and ventral \ lews of the holot> pe (L SNM 880085), off N'ietnam, South China Sea. Ma.ximum diameter = 51.1 mm. Type material: Holotype, USNM 880085, (max di- ameter 51.1 mm, min, diameter 44,7 mm, heiglit 43.8 mm), off N'ietnam, Soutfi China Sea, Paratype 1, An- seeuw collection, (max, diameter 68.0 mm, min. diam- eter 59.4 mm, height 60,4 mm), from the t\pe localits. Paratype 2, Dan Collection, (specimen damaged, max. diameter unknown, min. diameter 59.8 mm, height 56.5 mm) New Zealand C). Etymolof^y: This .species is named tor Dr. Bernard Me- tivier of the Laboratoire dc Malacologie, Muscimi na- tional d'Histoire naturelle, Paris, in recognition of his extensive work in the field of Malacology and his con- tributions to the knowledge of western Pacific Pleuro- tomariidae. Comparative remarks: Pciotrochus metivien is readi- ly distinguished trom all other species hitherto reported from the South C;hina Sea; Perotruchus hirasei Pilsbry, 1903 (Bondarev, 1991 ), F, teramachii kuroda,1955 (Lan, 1994; Okutani, 1979), P. caledonicus Bouchet and Me- livier, 1982, and /'. vicdani Kosuge, 1980 (Ravbaudi, p. Anseeuw and Y. Goto, 1995 Page 119 1992). In silhouette it most closeK resembles the southern Atlantic species Perotrochus atlanticus Rios and Ma- thews, 1968, particularly in the profile of the spire, but is otherwise unmistakeable. Of the other species from the western Pacific, P. tangaroana Bouchet and Metivier, 1982, is similar in size and construction and similarly lacks a clear color pattern, but can be distinguished from P. metivieri by its higher, less gradate spire profile, more inflated base with correspondingly rounder aperture, and shorter, narrower slit. Perotrochus nictivieri might be confused with juvenile P. teramachii Kuroda, 1955, but the former is smaller in adult form, lacks any clear pat- terning, and has a flatter base than P. teramachii. Per- otrochus metivieri resembles in some respects certain specimens of P. vicdani Kosuge, 1980, from the South China Sea, but these difter in having strong coloration and patterning as well as finely beaded sculpture on the teleoconch. Paratype 1 of P. metivieri differs somewhat in shape and appearance from the holotype, in particular by hav- ing a distinctly constricted penultimate whorl and slight- ly taller profile. Constriction of the whorl in mature adult specimens is a phenomenon occasionally observed in oth- er pleurotomariids and may be indicative of a gerontic stage. furnished the second paratype as well as valuable advice. The photographs were supplied by Mr. Paul Callomon of Elle Scientific Publications, Osaka, Japan, who also edited and prepared the manuscript. Finally, we thank two anonymous referees for their helpful comments. LITERATURE CITED Bayer, F. M. 1965. New pleurotomariid gastropods from the western Atlantic, with a summary of Recent species. Bul- letin of Marine Science 15(4):7.37-796. Bondarev, I. 1991, A rare shell in a fantastic seascape. Ha- waiian Shell News 39{7):.3, Lan, T. C. 1994. Correct locality data on some important shells originating from Taiwan, World Shells 9:112-114. Okutani, T. 1979. A new pleurotomariid gastropod from the Pacific Waters off Honshu, Japan. Venus 38(3);162-163. Raybaudi, L. M 1992 Pleurotomaria vicdani. World Shells 1;84. Rios, E. de C. and H, R. Mathews. 1968, Nova especie de Pleurotomariidae do Brasil (Mollusca: Gastropoda) Arq. Estaao de Biologia Marinha Universidade Federal do Ceara 8(l);65-68. Wagner, H P. and H. E Coomans 1990. Review of the Perotrochus africanus - complex, with a note on the no- menclature of the Western Australian species. Gloria Maris 29(3):41-52. ACKNOWLEDGEMENTS The authors are indebted to Mr. G. Campanini for bring- ing the first specimen to their attention. Mr. Donald Dan THE NAUTILUS 109(4):120-126, 1995 Page 120 Morphology and Possible Relationships of Ecphora (Cenozoic Gastropoda: Muricidae) Geerat J. \ ernieij Department of Geology and Center for Population Biology University of California at Davis Davis, CA 95616 USA ABSTRACT The muricid gastropod genus Ecphora Conrad, 1843, comprises Oligocene to Pliocene species from eastern North America and Oligocene to Middle Miocene species from Europe. Earlier workers speculated that it evolved in North America from an Early Oligocene Triionopsis-\\ke ancestor. Evidence from pre- viously unstudied shell characters indicates that Ecphora be- longs to the subfamily Ocenebrinae, whereas Tritonopsis is a subgenus of Cijmia Morch, 1860, a member of the Rapaninae. Ecphora lacks the parietal rib. Urate outer lip, and adapical extension of the outer lip of Cijmia and related genera, and may have been derived from an as yet unknown ocenebrine ancestor in which the external shell sculpture was predomi- nantly or exclusively spiral. Key icords: Ecphora. Stcnomphalus, Muricidae, Neogene, .At- lantic coastal plain. INTRODUCTION The genus Ecphora as ciassicalK understood by Amer- ican authors is a distinctive group that was characteristic of Oligocene to Pliocene faunas of the Atlantic Coastal Plain of the United States (Petuch, 1988a,b; Ward, 1992). Its status as a muricid is well established by virtue of its shell morphology and by the fact that very large muricid- type drill-holes are common in bivalves co-occurring vvitli Ecphora. These cylindrical holes are too large to have been made by muricids other than Ecphora (N'ermeij, 1987; Petuch,'l988b; Campbell, 1993). No consensus has emerged, however, about the position of Ecphora and its relatives within the Muricidae. Petuch ( 1988a, b) re- garded the group as a distinct subfamiK F.cphorinae, which he held to be a New World parallel to the Old World subfamily Rapaninae Gray, 1853. He circum- scribed the latter subfamiK narrowly around the geinis Rapana. Kool (1993b) tentatively assigned Ecphora to the Ocenebrinae, whereas Carter et al. (1994) suggested that it is deri\ed from Tritonopsis, a taxon assigned b\ all students of the Muricidae to the Rapaninae as broadK redefined b\ Kool 1 1993b), tfiat is, encompassing sucli genera as Thais. Rapana. Morula. Sassa. and their rel- atives. The purpose of this paper is to re\iew the genus Ecphora, to discuss the distribution within Muricidae of several shell characters that have not previously been considered in arguments about the taxonomic assignment of Ecphora and superficialK similar genera, and to argue that Ecphora should be referred to the Ocenebrinae. CRITICAL REVIEW OF PERTINENT GENERA Ecphora Conrad, 1843 Type species: Fusits quadricostatus Sa\-, 1824 (figures 1, 2). Synonyms: Stcnomphalus Sandberger, 1861 (type species, Fustis cancel- latus Thoma, 1845, non J. de C, Sowerby, 1826, = Slen- orr^phaltis caerulea (Romer-Biichner, 1827)) Chcsathais Petuch, 1988a (type species, Chesathais lindae Pe- tuch, 1988a). Ecphorosycon Petuch, 1988a (tspe species, Ecphora pamlico Wilson, 1987), Trisecphora Petuch, 1988a (type species, Ecphora tricostata Martin, 1904), Siphoecphora Petuch, 1988b (type species, Ecphora {Sten- omphaltts] aurora Wilson, 1987). Globccphora Petuch, 1994 (tvpe species, Ecphora floridana Petuch, 1988b). Latecphora Petuch, 1994 (t\ pe species, Ecphora hradtcyac Pe- tuch, 1987). Revised diagnosis: Protoconch multispiral; teleoconch broadK fusiform; suture deep, channeled in adtilt; false umbilicus present; sculpture consisting ot spiral elements onK, three to eight prominent cords on last whorl and two on preceding whorl; outer lip planar, thin, crenate at edge, smooth on apertural side; adapical junction of outer lip gratlual, \\ ilhout notch or extension; columellar margin smooth, straight; parietal rib absent, most species with thick brow II external calcitic la\er. I{emark>: Ecphora forms a compact group that, be- cause ot \ ariations in sculpture and shape, has been di- \ided into a number of subgenera by Petuch (1988a,b, 1994). These subgroups are here treated as subjective jimior synon\ms of Ecphora. The t\ pe species, Ecphora qiiadricostata. from the Pliocene Yorktown Formation G J. Vermel], 1995 Page 121 Figures 1. 2. Ecphora quadricostata (Say, 1824). Yorktowii Formation (Pliocene). Virginia; collected by P. VV. Signor; Vermeij collection. Ventral (1) and dorsal (2) views; scale bar = 1 cm. Figures 3, 4. Cymia vaughani (Mansfield. 1937). Haywood Landing Member. Belgrade Formation (latest Oligocene), North Carolina. E. H, \'okes Collection. Ventral (3) and dorsal (4) views; scale bar = 1 cm. of N'irginia (see Campbell, 1993), is characterized by four strong primary cords, and lacks secondary spiral sculp- ture. Most other species of Ecphora as defined here pos- sess secondary cords in addition to the primaries, which number from three to eight depending on the population or even the stage of ontogenv . Primary ribs vary in cross- section from triangular to T-shaped, as well as in width. Other variable characters are the size of the false um- bilicus and the length of the anterior siphonal canal. Carter et al. (1994) noted that all species of Ecphora beginning with the Late Oligocene £. tarnpaen.^is (Dall, 1915) have an external layer of calcite. The only fully aragonitic species is £. wheeleri, Rossbach and Carter, 1991 (Early Oligocene River Bend Formation, North Carolina), the earliest member of the genus in North .\merica. Petuch's tl9SSa) genus Chcsathais was based on his new species C. lindac from the Middle Miocene (Ser- ravalHan) Choptank Formation of Maryland. Petuch (1988a) cliaracterized this group as having a biconic fu- siform shell, lirate outer lip, and a sculpture of fine spiral ribs and threads. This description accords better with Rapana vaughani Mansfield, 1937 (figures 3,4) and R. biconica Dall. 1915 than with the ty pe species C. lindae. Petuch (1988b) included R. biconica in Chcsathais. but made R. rauglmni the type of Rapanecphora. which I consider to be a junior subjective synony m of Tritonopsis Conrad, 1865. As Ward (1992) already pointed out, Chcsathais lindac is a true Ecphora. The type specimen (L SNM 424254) has four primary spiral ribs on the last whorl and two on the penultimate whorl. The second cord, counting from the suture, forms the periphery, v\ hich is evenly rounded, .\ secondarv cord is situated below the suture. as well as between cords one to three and in front of the fourth cord. The whole surface, including that of the broad siphonal canal, is covered with fine threads. The columella is smooth and straight. There is neither a pa- rietal rib nor an anal notch, and the outer lip is not adapically extended. The inner surface of the outer lip is coarsely ribbed in accordance with the primary and secondary cords on the shells exterior. The false umbil- icus is very narrow. Chcsathais is therefore a junior sub- jective synonym of Ecphora. Whether its type species, Ecphora lindae, is a subjective junior synonym of E. n^eganac n}eganac Ward and Gilinsky, 1988, as Ward (1992) contends, must await studies of variation and com- parisons of material from the Choptank and Calvert For- mations of Mar> land. Until 1987, the name Stenon^phahis was used for Oli- gocene and Miocene species from Europe, Besides the type species, these include S. jaubcrti (Grateloup, 1840) from the Burdigalian (Early Miocene) and Helvetian ( = Langhian, Middle Miocene) of France; S. nwulinsi (Bro- chon, 1849) from the Burdigalian of France; S. wiech- rnanni (von Koenen, 1872) from the N'ierland Stage (Ear- ly Miocene) of the North Sea Basin; and S. koeneni (Gorges. 1952) from the Chattian (Late Oligocene) of Germany. Through the kindness of P. Lozouet, I have examined a specimen of Stcnon}phalus moutinsi from Saucats, France. The broadly fusiform shell has six spiral cords on the last w horl and two on the penultimate w horl. The second cord forms the obtusely angulate periphery , One or more secondary cords and threads of various sizes are situated between the primary cords. The subsutural ramp is vertical. The columella is straight and smooth. The outer lip is smooth on its apertural side, and lacks an anal Page 122 THE NAUTILUS, Vol. 109, No. 4 notch and an adapical extension. The long siphonal canal is covered with spiral tiireads. There is a narrow ialse umbilicus. The posterior end of the aperture lacks a parietal rib or knob. Stenomphalus jauberti is said to be similar, but has only three cords on the last whorl and shows partial separation of the adult last \\ horl from the penultimate v\ horl (Cossmann & Peyrot, 1924; Wilson, 1987). I have also e.xamined a lot labeled "Stenomphalus sp." in the National Museum of Natural History, Leiden, kindly loaned to me b\ A. W. Janssen. This lot is of unknown (but probably Early Miocene) age, and comes from a gravel quarry on the Resebergweg, Liibeck-Kiick- nitz, in Schleswig-Holstein, Germany. Of the six shells, four specimens have four primary ribs, one has three primary ribs, and one has five. Small secondary cords are observable anterior to the primary cords on some specimens. Two primary cords appear on the penulti- mate whorl. The periphery is marked by the second cord. As in .American species of Ecphora, the subsutural ramp is vertical, and there is a brown external layer probably composed of calcite. Cossmann and Peyrot (1924) referred the French spe- cies to Ecphora, which they considered a subgenus of Rapana. Wilson (1987) maintained this placement, but regarded Ecphora as a genus and placed the European species in the subgenus Steiiomphalus . He added a new American species, £. (S.) aurora, from the Pungo River Formation (Langhian Miocene) of North Carolina. Pe- tuch (1988b) made this species the type of his new sub- genus Siphoecphora. This species has a spindle-shaped shell with three very broad grooved spiral ribs on the last whorl, separated by narrow interspaces sculptured by one of two secondary spiral threads. As in the Eu- ropean species, two of the primary cords are visible in the spire whorls, and the subsutural ramp is nearly ver- tical. The species that have been assigned to Stenomphalus vary in shape and in the number of ribs, and I can fiiul no features that consistently distinguish Stenomphalus and Siphoecphora from Ecphora. Petuch's (1988b) pro- posal of Siphoecphora lor aurora was supported largely by the mistaken belief that the European species of Stenomphalus are small (less than 20 mm long). Al- though E. aurora (length 80 mm) is the largest species assigned to Stenomphalus, the Miocene species of that group from Europe are 50 to 75 mm long. As the name implies, species of Stenomphalus generally have a small false umbilicus, but so do many American species of undoubtwl Ecphora, especially the older si^ecies. I con- sider Stenomphalus and Siphoecphora to be junior sub- jective synonyms of Ecphora. Rapana Schumacher, 1817 Type species: Rapana foliacea Schumacher, 1817 ( = Ruccinurn hezoar Linnaeus, 1758) Diagnosis (shell characters only): I'loldcuiKli iiiultis- piral; teleoconch large (adults greater than 100 intn long). broadK lusiform; sculpture consists of scaly tuberculated spiral cords, seven or eight on the last whorl, between which are situated numerous fine spiral threads; suture deep; outer lip planar, thin, crenate at edge, lirate on apertural side, adapically extended, without anal notch; anterior to the fifth spiral cord is a spiral groove, w hich at the outer lip ends in three enlarged and ventrally directed crenations; columella straight, smooth; parietal rib absent; small false umbilicus present; thin external layer of calcite. Remarks: This genus is here narrowly circumscribed to contain three Recent species and their immediate Plio- Pleistocene ancestors. Besides the type species fi. bezoar, which occurs subtidally from Japan to the Philippines, I include R. venosa (Valenciennes, 1846) from the cool- temperate northwestern Pacific, and R. rapiformis (Born, 1778) from the Philippines and North Australia to the Red Sea. Rapana venosa is a recent immigrant to the Mediterranean and Black Seas. Other fossil species that have been assigned to Rapana belong to Cijmia and Ecphora. The Early Miocene Cyalifornian R. vaquero- sensis Arnold, 1907, and the Baja Californian Early Mio- cene R. imperialis Hertlein and Jordan, 1927, may be large ocenebrines related to Forreria and Chorus. Cymia (Cymia) Morch, 1860 Type species: Cuma sulcata Swainson, 1840 (= Buc- cinum tectum Wood, 1828) Diagnosis (shell characters only): Teleoconch thick, bi- conic; sculpture consisting of many fine scaly spiral threads and a single centrally placed spiral row of tubercles; aperture narrow; outer lip planar, thin, crenate at edge, coarsely lirate on apertural side, with deep adapical notch; columella straight, with strong central fold; parietal rib strong; suture indistinct except at final growth stage, when the last whorl may separate from the preceding one; false umbilicus narrow or absent; thin external layer of calcite present. Cymia (Tritonopsis) Conrad, 1865 Type species: Triton subalicatum C^onrad, 1848 Synonyms: Rapanccphora Petuch, 1988b (,tvpe species, Rapana vaughani Mansfield, 1937). Diagnosis: Shell fusiform to biconic; whorls flattened, separated by appressed or canaliculate suture; sculpture consisting of five or more spiral cords and secondary spiral threads; narrow false umbilicus present; aperture relatively broad; outer lip thin, lirate on apertural side, crenate at edge, with adapical extension hut w ithout anal notch; columella straight, with central fold that ma\ be obsolete in some species; parietal rib present, calcite layer absent Remarks: N'okes (1989) pouitcd out that the lineage leading to Cymia Iccia, « hich ll\cs in the eastern Pacific G. J. Vermei], 1995 Page 123 from Costa Rica to Peru, extends back to tlic latest Oli- goceiie or earliest Miocene of Peru us C. bcnyi Olsson. 1931). Other species are known from the Miocene of Trinidad, N'enezuela, Colombia. Panama, and Baja Cal- ifornia Sur, and from the Pliocene to the Recent of the tropical eastern Pacific. A second group of species that N'okes (1989) recognizes as belonging to Cyinia consists of species that differ from Cymio s. s. b\' having two or more rows of tubercles on the last whorl. These tubercles are situated behind the niidtlle of the last w liorl. and the subsutural slope is nearK vertical rather than obliciueK sloping as in the Cijinia tccta group. The shell therefore superficialK resembles double-knobbed specimens of the rapanine genus Stranionitu Schumacher, 1817. This group contains C. moiwplex (Sandberger, 1861) from the Stampian (= Ru- pelian, Earh Oligocene) of western Europe; C. gratc- loupi (d Orbigny, 1852) from the Aquitanian (latest Oli- gocene) of France; C. calcarata (Grateloup, 1833) from the Aquitanian and Burdigalian of France; C. hcnckcni Maur\, 1917, from the Early Miocene Baitoa and Tho- monde Formations of Hispaniola; C. marcanoi Yokes, 1989, from the latest Miocene or earl\ Pliocene Cercado Formation of the Dominican Republic; and perhaps C. picuriplicata Cossmann and Pe\ rot, 1924, from the Bur- digalian of France. The anal notch in this group is w eak. The outer lip is provided on its apertural side with den- ticles. Cords are numerous in the American species and in C. picuriplicata. but the\ number from four to si.x in C monoplcx. C. graicloupi, and C. calcarata. The cen- tral columellar fold is alwa> s strong. In C. calcarata there are one or two smaller columellar folds immediately anterior to the central fold. .\ thick parietal rib is in- \ ariabK present. This group of species ma\ deserve sep- arate subgeneric status. It differs from Cijmia s.s. by the presence of at least two posteriori)' placed rows of tu- bercles instead of one central row . The t\pe species of Tritonop.si.s. Triton i,ubahcatuiu Conrad, 1848, from the Mint Spring and B\ram For- mations of the N'icksburg Group (EarK Oligocene) of Mississippi, differs from the two groups of Cymia dis- cussed above b\ lacking an anal notch and b> the absence of tubercles. In some individuals, tw o of the spiral cords are more strongK expressed than the others (MacNeil & Dockery, 1984; Carter ct at.. 1994). The last whorl is rounded, not shouldered. The outer lip bears nine den- ticles on its apertural side that extend as lirae further into the aperture. A second species is C. (T.) woodii Gabb, 1860, from the EarK Miocene Kirkwood For- mation of New Jersey. A third undescribed species w ith an angulate rather than rounded last whorl and with axial swellings between the spiral bands was recorded b\ Woodring (1973) from the Gatunciilo Formation (Late Eocene) of Panama. The genus Rapanccphora Petuch, 1988b w as proposed for biconic shells with a low spire, sloping shoulder, and a sculpture of spiral threads and w eak peripheral nodes. Petuch (198Sb) based his genus on Rapana vaiighani Mansfield, 1937, from the Tampa Limestone (Aquitani- an) of Florida. Mansfield (1937) described this species on the basis of a cast made from a mold. OnK the upper surface of most of the last vvliorl and part of the spire is visible on the type specimen (USNM 495946). Characters of the aperture therefore remain unknow n. The last whorl bears eight or nine w eak spiral cords, of w hich the third (counting from the itulistinct appressed suture) forms a weakK noded periphery. Threads are intercalated be- tween the cords. Specimens from the coeval Haywood Landing Mem- ber of the Belgrade F'ormation of North Carolina have also been assigned to this species under the generic name Tritonopsi.s (Rossbach is Carter, 1991; Ward, 1992; Car- ter ('/ ai. 1994). I have examined beautifully preserved shells from this unit in the E. H. \'okes collection, as well as material sent to me under the name Tritonopsis laughani by J. G. Carter. Most specimens have five pri- mary spiral cords on the last whorl, but a few have six or seven cords. The third cord forms a smooth periphery, v.hich is located well posterior of the midpoint of the last w liorl. Between the primars cords are situated many spiral threads and sometimes a very weak secondary cord. The obliqueK' upturned anterior siphonal canal bears fine spiral threads. .Adjacent to it is a very small false umbilicus, w hich in main specimens is absent. The apertural side of the outer lip bears lirae or sometimes paired elongate denticles. The straight columella is usu- ally smooth, but in some specimens (Figure 2) a trace of a central fold is obser\able. The suture is narrowly can- aliculate. The outer lip is adapically elongated opposite a thick parietal rib, an anal notch is lacking. According to Carter et al. (1994), the shell is wholK aragonitic. Richards (1943) described Rapana gillcttci from the SiKerdale Beds (= Ha>wood Landing Member of the Belgrade Formation) of North Carolina. Ward (1992) regarded this taxon as distinct from his Tritonopsis vaitghani from the same formation. According to Ward, R gillcttci has weaker spiral ribs and more rounded whorls. I cannot separate R. gillcttci from shells consid- ered as T. vaiighani by Ward (1992) and Carter et al. (1994). Given the poor preservation of the type of Rapana vaughani and the tendenc\' for spiral sculpture to vary in prominence in many rapanines, I consider Rapana gillcttci Richards, 1943, to be a junior subjecti\e s\n- onym of Rapana vaughani Mansfield, 1937, w hich I con- sider to be a species of Cymia (Tritonopsis). Co-occurring with C. vaughani in the Tampa Lime- stone as well as in the Ha\ wood Landing Member of the Belgrade Formation is Rapana conica Dall, 1915. \\ard (1992) assigned this species to Tritonopsis. and pointed out that Richards (1943) referred to it as Rapana vaugh- ani. Petuch (1988b) assigned R. hiconica to Chcsathais. The species differs from Cymia (Tritonopsis) vaughani chiefl\ in having the spiral sculpture consisting of fine spiral threads, a sharp peripheral cord, and sometimes two to three additional cords below the periphery. The siphonal canal is longer and the columella is more curxed. No trace of a fold appears in the columella in any spec- imens I ha\e examined. Page 124 THE NAUTILUS, \'ol. 109, No. 4 Assignment of Rapana caughani and R. biconica to Tritoiwpsis (Rossbach & Carter, 1991; Ward, 1992; Car- ter et al.. 1994) is justified b\ virtue of the spiral sculp- ture, lirate outer lip, adapical extension of the outer lip, absence of an anal notch, and the presence in at least some specimens of R. vaughant of a very weak central columellar fold. Rapana conica clearly diverges signifi- cantU from the type of Tritonopsis in the absence of the columellar fold, but it is morphologicalK linked to the more t\pical members of the subgenus b\' R. caugh- ani. Yokes (1989) regarded Tritonopsis as a synonym of Cyniia. I belie\e that separation ol the two is warranted on the basis of the absence of an anal notch and of tubercles in Tritonopsis. The subgenus ranges from the Late Eocene to the Early Miocene of Atlantic North and tropical .America. Petuch (198Sb) included in his genus Rapanccphora one additional species besides Rapana vaughani. This is the Early Eocene R. nanjemoyensis Petuch, 1988b, from the Woodstock Member of the Nanjemoy Formation of Mar\ land. Carter et al. (1994) noted that the single spec- imen ot this species is poorly preserved. 1 have not seen it and cannot comment on its taxonomic placement. DISCUSSION Rossbach and Carter (1991) and Carter et al. (1994) proposed that Ecphora arose during the Early Oligocene from a TritonopsisAike ancestor. They pointed to a form of T. subaheata (Conrad, 1848) from the Byram For- mation of Mississippi with two strong cords on the spire whorls as close to the possible ancestor of Ecphora, whose earliest species (E. ivheeleri Rossbach & Carter, 1991) is known from the approximately coeval Ri\er Bend For- mation oi North Carolina. In support of this hypothesis, these authors noted similarities in spiral sculpture, shell shape, presence of a false umbilicus, and geographical proximity. Deri\ation of Ecphora from Tritonopsis would, how- ever, require the modification of four traits: the loss of the parietal rib, the loss of the central columellar fold, the loss of lirae on the apertural side ot the outer lip, and the elimination ot the adapical extension of the outer lip. The fact that each of these changes has occurred in one or another rapanine lineage shows that such a der- ivation is possible, but the alternative that Ecphora is unrelated to Cymia and Tritonopsis seems more parsi- monious to me. The reasoning in support oi this conclu- sion in based on the distribution in Muricitlae ot the io\n- traits enumerated above. The absence of a parietal rib appeals to be ple- siomorphic in the FaniiK Muricidae. This is the condition in all Trophoninae and Ocenebrinae, as well as in Rapana. The three oldest muricine genera (Paziella. I'oiricria, and Pterynottis). all of which existed b\ Paleocene lime, also lack a parietal rib. The rib is present in advanced niuricines such as Boliuus. Chicoreus. Uexaplex. Murex, I'hyllonotus, and Stratus, as well as in most rapanines. The only genera in the Rapaninae other than Rapana without a parietal rib are limpet-like taxa. These include Concholepas , in w hich the last whorl completely envel- ops all preceding w horls, and Plicupurpura. In the latter genus, only the limpet-like morplis of the two species, Plicopurpura columellaris (Lamarck, 1816) and P. patula (Linnaeus, 1758), lack the parietal rib (see Vermeij & Kool, 1994). Within Ocenebrinae. a fev\ species or pop- ulations have a parietal knob in the adult shell. This condition exists in Californian thick-shelled morphs of Niicella emarginata (Deshayes, 1839), in some individ- uals of the Californian Acanthinucella spirata (Blain- ville, 1832), and in the C^alifornian Roperia poulsoni (Carpenter, 1864) (see Vermeij, 1993). The absence of a parietal rib in Ecphora is consistent with assignment to the Ocenebrinae, and probabh represents the ple- siomorphic condition for the Muricidae rather than the secondary loss of a rib. The smooth columella is a character state that Ecphora has in common with most other muricids. Columellar folds are present in Rapaninae including Cymia. The absence of folds in Ecphora is probabK plesiomorphic in Ecphora and other Ocenebrinae, as well as in such rapanines as Rapana and Dicathais (for further discus- sion see Vermeij & Kool, 1994). Ponder and N'okes (1988) have similarK' argued that rugae on the columella are derived features in several muricine stocks such as Haus- tellutn and Bolinus. The apertural side of the outer lip of Ecphora is smooth. In Cyniia and many other Rapaninae, however, this sur- face is adorned with polished spiral ridges, or lirae Such lirae are never present in Ocenebrinae, although the apertural edge of the outer lip is often thickened and dentate. The plesiomorphic condition of Muricidae is probabK a smooth apertural surface of the outer lip. This is the condition seen in Ocenebrinae, Trophoninae, and in the plesiomorphic nunicine genera Paziella. Poirieria, and Ptcrynotus. The outer lip in Ecphora makes a high-angle junction with the preceding whorl, and is neither notched nor extended at its adapical end. In this respect it differs from Cymia and all other members of the Rapaninae except the limpet-like Concholepas. Ecphora shares its outer-lip junction with all other members oi the Ocene- brinae and Trophoninae, as well as with plesiomorphic Muricinae. Detailed anatomical studies bv Kool i, 1993a, b) have revealeil that fusiform nnnicids w ith a moderatelv long open siphonal canal and without well-defined varices belong to two distinct clades. the Ocenebrinae Cossmann, 1903, and the Rapaninae Grav, 1853. The Ocenebrinae in addition contains a large group of species with a nar- row, sometimes closed, siphonal canal and with predom- inant axial sculpture. In a subsequent studv (N'ermeij & Kool, 1994), we idenlitied shell characters by which the two subfamilies can be distinguished. .\s pointeil out above, almost all rapanines have a parietal rib, and the outer lip is either adapicallv extended or provided \\itli an anal notch. Ocenebrinae lack these traits. G. J. Vermeij, 1995 Page 125 Although a formal cladistic anaK sis of shell characters must be performed and is in progress to test these ar- guments, the distribution of characters implies that Ec- phora shows none of the synapomorphies associated with the Rapaninae (N'ermeij & Kooi, 1994), I therefore agree with Kooi (199.'3b) that Ecphora should be assigned to the Ocenebrinae. The only fact that potentially conflicts with this interpretation is the finding that Ecphora pam- lico Wilson, 19S7, and E. cf. E. quadricostata (Sa\, 1824) have a three-whorl (nuiltispiral) protoconch (Ward, 1992; Kooi, 1993b). Multispiral protoconchs occur widely among Rapanines and are apparently primitive in the Muricidae (Yokes, 1971; Ponder & Yokes, 1988). The protoconch of Ecphora differs from that of rapanines, however, h\ lacking a sinusigeral notch Most Ocenebrinae ha\e pau- cispiral protoconchs, but there are exceptions. Houart (1989) has noted that three West African species of Oce- nebra, (O. coseli Houart, 1989; O. inermicosta Yokes, 1964; and O. isaacsi Houart, 1984) have a multispiral protoconch. The same ma> be true of the New ZealantI ocenebrine genus Haustrum (Kooi, 1993b). Unfortunately, the origins of Ecphora within the Oce- nebrinae remain obscure. Wilson (1987) and Petuch (1988b) justifiably rejected the idea that the Late Cre- taceous species Ecphora proquadricostata (Wade, 1917) from the Riple\' Formation (Maastrichtian) of Tennessee is related to true Ecphora. On the basis of protoconch morpholog), Bandel (1993) argues that £. proquadri- costata is a mesogastropod related to the Trichotropidae. Other Cretaceous genera from the Gulf Coastal Plain assigned to the Muricidae b% Sohl ( 1964a, b) have also been interpreted bs Bandel (1993) as belonging to this mesogastropod famiK . There are no obvious Eocene an- tecedents of Ecphora in North .'\merica. Many ocenebrine genera share with Ecphora a sculp- ture consisting predominantK' or e.xclusiveK of spiral cords or heav\ ribs. These include the South .American genus Chorus (Miocene to Recent) and the Floridian Pliocene genus Zulloia (see Petuch, 1994), as well as members of the so-called Nucella group: Acanthina in temperate South America, Acanthinucella in the north- eastern Pacific, Lepsiella and Lcpsithais in .Australia ami New Zealand, Sucella in the temperate North Pacific and North Atlantic, and Trochia in South Africa. None of these genera typically possesses a false umbilicus as does Ecphora, and the outer lip of many members of the Sucella group is greatK thickened and denticulate on its apertural side. Most species of Ecphora have the outer lip thin and smooth on its inner side. This condition also occurs in the South African Trochia cingulata (Lin- naeus, 1767), the t>pe of Trochia Swainson. 1840, which further resembles Ecphora in having one to fi\e heav\ spiral ribs. None of the genera mentioned above is known from before latest Oligocene time, but a pre-Miocene origin of the Mucella group is strongly implied b\' the Recent distribution of its members (N'ermeij, 1993). Ec- phora is first known in the EarK' Oligocene of eastern North America and France. Where the group originated cannot be pinpointed with the available evidence. The longer persistence of Ecphora in the Western .At- lantic than in the European sector of the eastern .Atlantic is an example of a common biogeographical pattern in Neogene mollusks. Europe lost many genera after the Middle Miocene that survived in tropical America, the western Atlantic, of the Indo-West Pacific region. Pos- siblv the extinctions of the Middle Miocene were more severe in the eastern .Atlantic than elsewhere. Further discussion of this pattern will be deferred to a later paper. Petuch (1988a) proposed the new subfamily Ec- phorinae to encompass Ecphora and several other genera that have here been synonymized with Ecphora. He considered it to be a parallel development to the Old \N orld Rapaninae, narrowly defined around the genus Rapana. Here I consider tlie Ecphorinae to be a genus m the Ocenebrinae. Whether the morphological resem- blance between Rapaninae and spirally sculptured Ocenebrinae (including the Sucella group) is the result of con\ergence or of the retention of plesiomorphic fea- tures coumon to both groups remains an open question. ACKNOWLEDGMENTS I thank Pierre Lozouet (Museum National d'Histoire Na- turelle, Paris), A. W. Janssen (Nationaal Museum voor Natuurlijke Historic, Leiden) and Warren Blow (U.S. National Museum of Natural History, Washington) for the loan of specimens; Joseph G. Carter (Uni\ersity of North Carolina, Chapel Hill) for sending me specimens; Gary Rosenberg (.Academ\ of Natural Sciences of Phil- adelphia), EniiK N'okes (Tulane L iii\ersit> ), and David Dockery III (Mississippi Geological Survey, Jackson) for the opportunity to work with collections in their care; Janice Cooper, Edith Zipser, and Hermine Yermeij for technical assistance; and the National Science Founda- tion for funding this project. LITER.ATURE CITED Bandel, K. 1993. Caenogastropoda during Mesozoic times. Scripta Geologica, Special Issue 2:7-56. Campbell, L. D. 1993. Pliocene molluscs from the Yorktown and Chowan River Formations in \ irginia. \ irginia Di- vision of Mineral Resources Publication 127.1-259. Carter, J. C, T, J, Rossbach, K. J. Robertson and L. W. Ward. 1994. Morphological and microstruclural evidence for the origin and early evolution of Ecphora (Mollusca: Gastro- poda). Journal of Paleontology 68:905-907. Cossmann, M. and A. Peyrot. 1924 (1909-19:35). Concholo- gie neogenique de I'.-Vquitaine. .Actes de la Societe Linneenne de Bordeaux 63-86: 1-3600. Houart, R. 1989 Description of a new species of Ocenebra (Muricidae: Ocenebrinae) from western .Africa. Publica- ^oes Ocasionais da Sociedade Portuguesa Malacoiogica 13: 63-64. Kooi, S. P. 1993a. The systematic position of the genus S'ucella (Prosobranchia: Muricidae: Ocenebrinae). Nautilus 107: 43-57. Page 126 THE NAUTILUS, Vol. 109. No. 4 Kool, S. P. 1993b. Phylogenetic anaKsis of the Rapaninae (Neogastropoda: Muricidae). Malacologia 35:155-259. MacNeil, F. S and D T Dockerv HI. 1984. Lower Oligocene Gastropoda, Scaphopoda, and Cephalopoda of the N'icks- burg Group in Mississippi. Mississippi Department of Nat- ural Resources, Bureau of Geolog\, Bulletin 1241-415. Mansfield, W. C. 1937. Mollusks of the Tampa and Suwannee Limestones of Florida. Geological Bulletin (Tallahassee, Florida) no. 15:1-334. Petuch, E. J. 1988a. New species of Ecphora and ecphorine thaidids from the Miocene of Chesapeake Bay, Maryland, U.S..\. Bulletin of Paleomalacology 1:1-16. Petuch, E. J. 198Sb. Field Guide to the Ecphoras. Coastal Research Foundation, C^harlottesville, VA. Petuch, E. J. 1994. Atlas of Florida Fossil Shells {Pliocene and Pleistocene Marine Gastropods). Chicago Spectrum Press, Evanston, Illinois. Ponder, VV. F. and E. H. Yokes. 1988. A revision of the Indo- West Pacific fossil and Recent species of Murex s.s. and Haufitellum (Mollusca: Gastropoda: Muricidae). Records of the Australia Museum. Supplement 8:1-160. Richards, H. G. 1943. .■\dditions to the fauna of the Trent Marl in North Carolina. Journal of Paleontoiog\ 17:518- 526. Rossbach, T. J. and J. G. Carter. 1991. Molluscan biostratig- raphy of the lower River Bend Formation at the Martin Marietta Quarry, New Bern, North Carolina. Journal of Paleontology 65:80-118. Sohl, N. F. 1964a. Neogastropoda, Opisthobranchia and Basommatophora from the Ripley, 0\\ 1 C^reek, and Prairie Bluff Formation. U. S. Geological Sur\ey, Professional Pa- per 331-B:153-344. Sohl, N. F. 1964b Gastropods from the Coffee Sand (Upper Cretaceous) of Mississippi. U. S. Geological Survey, Pro- fessional Paper 331-C:345-394 N'ermeij, G, J. 1987. Exolution and Escalation: an Ecological History of Life. Princeton Lniversit) Press, Princeton, NJ. Vermeij, G. J. 1993. Spinucella, a new genus of Miocene to Pleistocene muricid gastropods from the eastern Atlantic. Contributions to Tertiary and Quaternarv Geologv 30:19- 27. X'ermeij, G. J. and S P. Kool. 1994. Evolution of labral spines in Acanthaif^. new genus, and other rapanine muricid gas- tropods. N'eliger 37:414-424. \'okes, E. H. 1971. The geologic history of the Muricinae and Ocenebrinae. Echo 4:37-54. Yokes, E. H. 1989. Neogene paleontology in the northern Dominican Republic 8. The family Muricidae (MoUusca: Gastropoda). Bulletins of .American Paleontolog) 97:5-94. Ward, L. \V. 1992. Molluscan biostratigraphy of the Miocene, Middle Atlantic Coastal Plain of North .America. X'irginia Museum of Natural Historv Memoir 2:1-159. Ward, L. W. and N. L. Gilinsky, 1988. Ecphora (Gastropoda: Muricidae) from the Chesapeake Group of Mar\land and Yirginia. Notulae Naturae 469:1-21. Wilson, D. 1987. Species of Ecphora, including the subgenus Stenoniphalus, in the Pungo River Formation. Smith- sonian Contributions to Paleobiolog) 61:21-27. Woodring, W. P. 1973. Geology and paleontolog) of Canal Zone and adjoining parts of Panama: description of Ter- tiar> mollusks (addditions to gastropods, scaphopods, pe- lecypods: Nuculidae to Malleidae). L .S. Geological Survey Professional Paper 306-E:439-539. THE NAUTILUS 109(4):127-1;3S, 1995 Page 127 Taxonomic Notes on the Western Atlantic Turridae (Gastropoda: Conoidea) Donn L. Tippeit 10281 Gainsborough Road Potomac, MD 20854, ISA ABSTRACT This paper presents results ol research on the systematics of the western Atlantic Turridae. The following new species are de- scribed: Drillia {Dhlliajuoljci, Dnllia [ClathrodriUia) pctuchi . Fenimorea kathijae, F. petiti, Hcdiliopsis riosi, and Claihiirella eversoni. The new name Drillia (ClathrodriUia) dautzcnbcrgi is proposed, as are the new generic combinations Fenimorea pagodula and Pilsbryspira (Syniphispira) aubcrti. Lectotypes are designated for Drillia {ClathrodriUia) dautzcnbcrgi and Fenimorea pagodula. Radulae are figured for Drillia (Drillia) wolfei. Lioglyphostoma hendersoni, ViridrUlia uilliami. V. hendersoni, Inodrillia nucleata, Pilsbryspira albocincta, Pyr- gocythara filosa. P. plicosa and P. danae (U. S. west coast). Opercula are figured for: Drillia (Drillia) wolfei, Fenimorea kathyae. F. sunderlandi var., F. petiti and Lioglyphostoma hendersoni. The animal of Pilsbryspira albocincta is figured. Key Words: Turridae. Ta.xonomy, Western Atlantic. INTRODUCTION Although the family Turridae in the western Atlantic has been extensively studied and documented, there re- main many gaps in our knowledge of its systematics and a major review is needed. New species are discovered not infrequently . In the absence of a definitive revision, such as provided for the tropical eastern Pacific by Mc- Lean (1971) and currently being carried out bv Kilburn (1983, 1985, 1986, 1989, 1991, 1992, 1993, 1994) for the South African fauna, scattered information is at least of some value. This paper presents findings obtained during research on the family conducted over a number of years. Included are descriptions of new species, a new name, lectoty pe designations, animal and radular details, range data, and other information. It is hoped that these find- ings will provide clarification of some issues and repre- sent a contribution to our knowledge of the family. MATERIAL AND METHODS Specimens, both of empty shells and shells containing either preserved or dried animals, were obtained from various sources. Dried animal material was dissolved in KOH Radulae were mounted on microscope slides and stained with Prontocil + CMCP-10. Type specimens, slides, and voucher specimens were deposited at the I'SNM and other institutions. The conventional classifi- cation of the family Turridae is used rather than that proposed by Taylor et ai. 1993. This proposed classifi- cation is too recent to have gained general acceptance and does not consider the fossil record of the family as noted in a re\iew of the classification h\ Kohn and Mc- Lean (1994). Abbreviations used: a = ratio of length of aperture plus canal to shell length; N = nimiber, (of adult specimens); S.D. = standard deviation; w = ratio of maximum shell w idth to length; AMNH = American Museum of Natural Historw New- York; ANSP = .\cademy of Natural Sciences, Philadelphia; DMNH = Delaware Museum of Natural History, Wil- mington; IRSNB = Insitut Ro\al des Sciences Naturelles de Bel- gique, Brussels; LACM = Los Angeles County Museum of Natural His- tory, Los Angeles; MCZ = Museimi of Comparative Zoology, Harvard I'ni- versity , Cambridge; MNHN = Museum National d'Histoire Naturelle, Paris; MORG = Museu Oceanografico "Prof Eliezer de C. Rios , Rio Grande; NHM = The Natural History Museum, London; NM = Natal Museum, Pietermaritzburg; USNM = National Museum of Natural History, Smith- sonian Institution, Washington, DC. SYSTEMATICS Subfamily Drilliinae Olsson, 1964 Genus Drillia Gray, 1838 Type species: Drillia uinbilicata Gray, 1838; by sub- sequent designation (Gray, 1847). Drillia (Drillia) uolfei Tippett, new species (Figures 1, 28, 31) Page 128 THE NAUTILUS, Vol. 109, No. 4 D. L. Tippett, 1995 Page 129 ? Sp "L" Rice 6; Kornicker. 1965: 129, pi. 8. &g. 6. Description: Shell small (14.1-19 mm, mean = 15.7, S.D. ± 1.4, N = 12), elongate-fusiform with tall spire, anterior canal of moderate length, emarginateil parie- tally by strong callus but lacking umbilical chink (w = 0.33-0'38, a = 0.34-0.38). Protoconch of two smooth whorls, nuclear tip laterally placed. Teleoconch whorls initialK flat-sided, with slightK concave shoulder slope on last whorl. .\.\ial sculpture of wideK spaced, rounded ribs, about 10 per v\ horl, extending from suture to suture on early w horls, sinuous, reduced in strength on shoulder slope of later whorls. Spiral sculpture of slightK wavy. e\enl\ spaced striae. Sinus on shoulder slope, deep, I -shaped, rather narrow, bordered medialK b\ a pari- etal tubercle that does not constrict opening. Outer lip thin, sharp, with small but distinct stromboid notch at lower e.xtremitv'. \'ari.\-like enlarged axial rib about 1/ 4 whorl behind outer lip. C^olor uniform light brown Operculum (figure 31) leaf-shaped with terminal nucle- us. Radula (figure 28) drilliine, with numerous rows of teeth, each consisting of small, unicuspid rachidian tooth, two comb-like lateral teeth, and two long, sharp-pointed marginal teeth. Dried animal, as viewed during treat- ment w ith KOH, with prominent eyes on short e\ e stalks, mantle with strong anterior siphonal extension and dis- tinct, curved indentation on the right for anal sinus. Types: Holotvpe. USNM 880070. Paratvpes (donated bv Dr. Wolfe), one each at AMNH, MCZ, DMNH, ANSP, LACM, \M, MORG. \HM. MXHX. IRSXB, author s collection, others in the Wolfe collection. Type locality: 15 km east of Cape Lookout. North Car- olina at 20 fm (36 m), from the Elmer Dewey Willis scallop plant, Williston, North Carolina. Leg. Dr. Doug- las Wolfe, March-Ma\ 1971. The North Carolina scallop fisher\ at the time of col- lection of the present material was concentrating on an area approximately 15 km east of Cape Lookout in about 20 fathoms according to Schwartz and Porter (1977). This is therefore selected as the t\pe localit\. SubsecjuentK this scallop plant began to shuck scallops imported from off central to northern Florida. Despite numerous col- lecting excursions at various scallop shucking houses in Carteret County, North Carolina, both before and after the above dates, this species was not found again. Range: Off North Carolina; ? Campeche Bank, Yu- catan. Remarks: Drillia iiolfei superficially resembles Feni- morea pagodtila Dall, 1889 (figure 3), It differs in ha\ing narrower, fewer, and more wideK spaced ribs that do not run from suture to suture, stronger and more widely separated spiral threads that are not of the "butterfly wing t\pe (see below), a weaker varix. and in lacking a spiral color band on the whorl peripher\ , Despite the geographic separation, Rice and Kornicker's (1965) Sp. "L" (figure 2), from the Campeche Bank off Yucatan, appears to be this species. There is a high percentage (about 1/3) of shells with repaired damage, probably indicating attempts at predation b>' crabs. Named for Dr. Wolfe for his contribution of the material and assis- tance in developing important data on collection locality. Drillia (aathwdrillia) Dall, 1918 Type species (of subgenus): Pleuwfoma gibbosa Reeve. 1843 (= Murcx gibbosus Born, 1778). Remarks: Clathwdrillia is conventionally regarded as a subgenus of Drillia {e.g.. McLean. 1971. Abbott. 1974. Kilburn. 1988. S\soev in Taylor et ai. 1993). Members of the subgenus have stronger spiral sculpture and a more pronounced terminal varix blotched with color than the nominal genus, Drillia (Clathrodrillia) dautzcnbcrgi Tippett. new name (Figures 4. 5) Drillia gibbosa var. minor Dautzenberg, 1900:20, pi 9, fig. 2 (anterior & posterior views), non Drillia minor Seguenza, 1880. Clathrodrillia minor (Dautzenberg, 1900) Rios, 1975:130, #580. pi 39; Altena, 1975:7, pi 7. figs. 1,2. Description: Shell medium sized (to 27 mm), elongate- ovate, with tall spire, moderately-sized body whorl ter- minating in barely differentiated, notched, recurved an- terior canal (w = 0.38, a = 0.33). Protoconch of two smooth whorls, with lateralK' placed, immersed tip, bear- ing two or three brephic axial ribs at termination of protoconch. Teleoconch of about 9 moderately rounded whorls, with strong sulcus on upper third Sculpture of numerous, regularK spaced, narrow, rounded axial ribs separated by equal interspaces and decussating, crowded spiral cords separated by deep grooves. Axial ribs reduced in strength, curved on sulcus, spirals cords finer, more on sulcus. Varix low, broad, about 1/4 whorl back from Figure 1, Drillia (Drillia) woljei Tippett, new species, Holot\pe, L'SNM 880070, 16.6 x 6.1 mm. Figure 2. Sp. "L" Rice & Kornicker, 1965, L'SNM 667703, 12.1 x 4.2 mm. Figure 3. Fenimorea pagodula (Dall, 1889). Lectotype. USNM 87471, 17.7 x 5.8 mm. Figure 4-5. Drillia {Clathrodrillia) dautzenbergi Tippett, new name. 4. Lectotype, IRSNB, 24,2 x 9. ,3 mm. 5. USNM 880076. 27.1 x 10.2 mm. Figures 6-7. Seddiopsis riosi Tippett. new species. 6. Holotvpe. MORG 31.775. 14,6 x 5.4 mm. 7. Ape.x, 12 X, Paratype, MORG 31,776. Figure 8 Sediliopsis gracilis (Conrad, 1830) Apex, after Gibson, 1962, pi. 41, fig. 11. Figure 9. Lioglyphostoma hcndcrsoni (Bartsch, 1934). USNM 880078, 13.7 x 5.4 mm. Figures 10- 11. Clathurella eversoni Tippett, new- species. Holotvpe. USNM 880074, 4.5 x 1,7 mm. 10. Anterior view. 1 1. SEM of aperture show ing teeth, 22 x , Figure 12. Fenimorea janetae (Bartsch, 19341. Holotvpe, L'SNM 430249, SEM showing "butterfl\ wing" microsculpture, 22 x. Page 130 THE NAL TILLS, \ol. 109, No. 4 outer lip. Lip fluted by spirals cords. Siphonal fascicle inoderateK de\ eloped. Sinus deep, U-shaped, occupying entire sulcus, with reflected rim terminating in parietal tubercle continuous with marginated cohunellar callus bearing weak false umbilicus below . Outer lip with broad, shallow stromboid notch just above anterior end. Color pure white. Type: Institut Ro\al des Sciences Xaturelles de Bel- gique, one specimen, no catalog number, here selected as lectot\pe. Type locality: Isla Margarita, Venezuela. Range: \\'idespread in tlie lower Caribbean and north Brazil. Remarks: Both Rios (1975) and Altena (1975) recog- nized that this species is not a h)rm of D. (C.) gibbosa as thought by Dautzenberg. It differs in being much smaller and in having more delicate sculpture, including finer spiral cords, and a weaker varix, siphonal fasciole and parietal tubercle. It is all white in color and lacks the color blotch on the vari.x that is present in D. (C.) gibbosa. Drillia dautzcnbcrgi has spirals on the sulcus, which are lacking in D. (C.) gibbosa. Both Rios (1975) and Altena (1975) not only distinguished this species from D. (C.) gibbosa. but also raised Clalliwdrillia to full generic rank, a step not taken in the classification herein. As a consequence of being retained in the genus Drillia, this species name is preoccupied by Drillia minor Seguenza, 1880 (Seguenza, 1880:103, pi. 11, fig. 8), a species from the Miocene of Italy. A new name thus becomes necessary and is supplied here. The species is named after the original author. The specimen from IRSNB (figure 4) is housed in a circular bo.x containing an old identification slip stating "Drillia gibbosa Born var. minor Dautz. Type. pi. A, f. 2.2, I. Margarita, 24 I 9(i" The same is written on the bottom of the container. A label of Gilbert's states the specimen to be the holotype. Comparison of the shell with Dautzenberg's original figures shows that it was undoubtedly used for these illustrations. The\ are iden- tical including repaired breakage preceding the lip and a somew hat sliort anterior canal. There is a second healed break about liali\\a\ through development of the body whorl that alters the continuity of the canal, making it shorter than normal. Altena's (1975) figures and the spec- imen shown here (figure 5) demonstrate the full devel- opment of the canal. The specimen illustrated in figure 5 has slightly finer a.xial and spiral sculpture and more and finer spirals on the sulcus than the IRSNB specimen, but otherwise shows the same characters. Dautzenberg did not designate a holotype. The species was originally reported as being from various South Caribbean locali- ties. Altena (1975) mentions ha\ing seen syntypes. As Gilbert's assignment v\as not published, it cannot be con- sidered a lectotype designation. As designation of a lec- totype is appropriate, this specimen [IRSNB (figure 4)] is here designated as the lectotype. Drillia (Ctathrodrillia) petiichi Tippett, new species (Figure 18) ? Crassispira sp. Sutt\, 1986:96, photo. Splaidrillia sp. Petuch, 1988:160, pi. 38, figs. 3,4. Description: Shell moderateK large (to 53 mm), fusi- form, turreted, with tall spire. Body whorl somewhat truncated, ending in short, open, slightly notched, re- cur\ed anterior canal (w = 0.39, a = 0.43). VN'horls ca. 13 including 2 smooth nuclear whorls with immersed tip. Teleoconch whorls rounded, with deepK concave shoulder slope. Periphery angulated along upper ends of axial ribs, situated 1/3 whorl below suture. Sculpture of strong, rounded axial ribs (about 14 on penultimate whorl) with equal interspaces, with fine spiral striae that are absent on sulcus. Massive varix 1/4-1/3 whorl behind flaring, fluted outer lip, bearing small but strong strombo- id notch just above end of anterior canal. Axial ribbing reduced or absent follow ing varix. Sinus deep, L -shaped, margined by recur\ed callus, bearing parietal tubercle, its entrance narrowed by upward extension of outer lip edge. Siphonal fasciole present. Large specimens with small umbilical chink. Color w hite, with 3 brow nish spi- ral bands, adapical band spotted, on rear surface of ribs. Varix shows a blotch of same color. Type: Holot\ pe, USNM 880071. Type locality: Off Barbados, 470 ft (142 m). crabbed, Dec. 1986, D. Hunt! Additional material examined: AMNH 186435, 1 spec; author s collection, 1 spec. Range: Known oiiK from Barbados. Remarks: Drillia petitchi is similar to Drillia (Clath- rodrUlia) gibbosa, but difiers in being more tumid and robust, having fewer and broader ribs, having a less re- curved but more strongly notched anterior canal, and in having a base color that is w hite rather than pale brown. The varix is much more strongK colored in D. (C.) gibbosa. which lacks the spiral banding. Figure 13. Fcninwrca jaiwlae (.Barlsch, 1934). Holotype. l^SNM 430249, 37.8 x 14,3 mm. Figure 14. Fcnimorca kathijac Tippett. new .species. Holotype, USNM 880072, 36.1 x 14 iiuii. Figure 15. Fcnimorea xundertandi (Petuch, 1987) Holot\pe, USNM 859800. 40 X 14 mm. Figure 16. Fenirnorea mndcrlandi (Petuch, 1987) var. USNM 880077, 52 x 17.9 mm. Figure 17. Fenimorea petiti Tippett, new species. Holotype, USNM 880073, 13.2 x 5.2 mm. Figure 18. Drillia (Clathrodrillia) petuchi Tippett, new species. Holotype, L'SNM 880071, 39.4 x 15.4 mm. Figure 19. Pilsbryspira (Niimphispira) auberti (Lamv, 1934), new combination. USNM 880079, 34.7 x 12,8 mm. D. L. Tippett, 1995 Page 131 Page 132 THE NAUTILUS, Vol. 109, No. 4 Figures 20-28. Radular teetli. Scale bars = 100 ^m 20. In- odrillia nucleata. L'SNM 874.54. 21. Viridrillia williami. USNM .319167, 22. Viridrillia hendersorn. L'SNM .3.54327. 23. Pyr- gocythara plicosa. USNM 880083 (specimen sacrificed). 24. Pyrgocythara danae. L'SNM 266.350, 25. Pyrgocylhara filosa. L'SNM 880082 (specimen sacrificed), 26. Lioglyphostoma hen- dersoni. USNM 880080 (specimen sacrificed). 27. Pilsbryspira albocincta. USNM 880081 (specimen sacrificed), 28. Drillia wolfei. USNM 880084 (specimen sacrificed). Figure 29. Pils- bryxpira albocincta. USNM 880081, Schematic drawing of ani- mal, anterior area opened: Figures 30-34. Opercula 30. Feni- morea swulerlandi var, USNM 880077. 10,0 x 5.5 mm, 31. Drillia uolfei. L'SNM 880070, 3,4 x 1,7 mm, 32. Fcnimorea kathyac. L'SNM 880072, 4.5 x 2,6 mm, 33. Fcnimorea petiti. L SNM 880073, 2,4 x 1.3 mm, 34. Lioglyphostoma hender- soni. USNM 880078, 3.0 x 1,5 mm, ct, ctenidium; es, eye stalk; f, foot; m, mantle edge; pb, pro- boscis; rhc, rhynchodeal cavity; rhr, rhynchodeal roof, reflected; si, siphon. PetLicli (1988:160) considered this species to be an example of his Barbadan Secondary Relict Pocket forms and a po.ssible relative of the Splendrillia scala {=Drillia .s(Yi/(; I'ilsbr\ & Johnson, 1917:15.5) complex of his Gura- ban snliprovince. However Splendrillia scala is not a Clathrodrillia. whereas the present form is a typical member of the group having the characteristic hump- backed form blotched with col.-r, as well as distinct, relatively coarse, spiral threading. Barbados, the type locality of D. (C.) peluchi, is near the \Cnezuelan sub- province, where D. (C.) gibhosa occurs. A more similar form is Jung"s (1965:567) Clathrodrillia ?n. sp. aff. C. islaliiulac i\Iaur\ , 1917) In mi the Miuccnc of the Parag- uana Peninsula, Venezuela. That species is a typical clathrodrilliine, has the same number of axial ribs as D. {Opctuchi. but is narrower and the axials tend to extend sinuousK over the sulcus to the suture above. It is possible that Maurv's species is an ancestor of both D. (C. ) gibbosa and D. (C.) peluchi. The present species is named for Dr. Edward J. Petuch, who first noted the species, and who has made significant contributions to the Caribbean molluscan fauna. Genus Fcnimorea Bartsch, 1934 Type species: Fcnimorea janetac Bartsch, 1934. Remarks: Bartsch created the genus Fenimorea for his new species F. janetac on the basis of its odd micro- sculpture of fine axial and spiral striae that produces a characteristic wavy pattern that "resembles the scales on some butterfly wings ' (Bartsch, 1934:3. See figure 12). He was aware that there are other species with the same characteristic, including Dall's F. moseri (Dall, 1889) and F. jiicata Reeve, 1845, but did not assign them to his new genus. Fenimorea kathtjae Tippett, new species (Figures 14, 32) Splendrillia sp. Petuch, 1988: pi 38, figs. 5,6. Description: Shell medium sized (to 36 mm), fusiform- biconic with moderately tall spire. Body w horl gradually narrow ing, terminating in scarceK differentiated, short, notched siphonal canal (vv = 0.39, a = 0.37), Whorls ca. 11 including protoconch of two smooth whorls. Teleo- conch whorls moderately rounded, with concave sulcus on upper 1/3, roundl\- angulated below by upper ends of axial ribs. Sculpture of strong, rounded axial ribs (about 10 on penultimate w horl) w itii narrow er interspaces. Spi- ral striae fine, wavy, foriuing 'butterflv wing" pattern in later whorls. Sinus deep, U-shaped, w ith parietal tu- bercle, entrance narrowed b\ an upward extension of outer lip. Constriction varies from slight to almost com- plete closure. Large \arix 1 3 whorl behind lip. Strombo- id notch shallow. Columellar callus variably emarginate. Color white, tvpicalK with three red spiral bands, the most prominent peripheral and \ isible on spire just abo\ e suture. Varix slightK blotched w ith red on forward sur- face. Operculum (figure 32) leaf-shaped with terminal nucleus. Type: Holot>pe. USNM 880072. Type locality: West coast Barbados, dredged, 500 ft (152 m), Oct 1986, D. Hunt! Additional material examined: 1 specimen, dredged at 300 ft (91 m). off Egniont Ke\ , FL; 1 specimen, Hy- poluxo, FL. 190 ft, (58 m), both in author's collection. Range: Barbados; peninsular Florida, Remarks: The present species is another example of the F, janetac complex of attractive shells. When fresh the shells are shinv' and colorful, luuing bright spiral D, L. Tippett, 1995 Page 133 banding in various shades of orange, red-brown or red. Fenimorea kathijac is most similar to F. janctae (figure 13), (w = 0.38, a = 0.36), having shouldered whorls and a sulcus but with a shorter spire, longer base, as well as fewer and broader ribs. The spiral banding in F. janctae is orange and broad but is bright retl and narrow in F. kathyae. F sundcrlandi (Petuch, 1987) (figure 15), (w = 0.35. a = 0,34), another member of this group, lacks a sulcus, has non-angulate whorls, and narrower, more numerous ribs that are opisthocline, rather than orthocline, as in F. janctae and F. kathyae. As noted b\ Petuch (1987:19), F. sunderlandi has been misidentified as F. janctae. The shells so misidentified (figure 16, figure 30 for operculum), (w = 0.34, a = 0.35), coming principalK from the well-known "bushels, dredgings. of Jim Moore and Riley Black in the 1960s and 1970s, are a \ariet\ of F. sunderlandi that is larger and narrower. This form has a facies suggestive of F. janctae. but shows the whorl- outline and rib characteristics of F. sunderlandi. All members of the group have the "butterfl\ wing spiral sculpture although it differs in strength from specimen to specimen. The species is named for another beaut\ , and fine field collector, the author's \\ ife. Fenimorea petiti Tippett, new species (Figures 17, 33) ? Drillia albicoma Dall. 1889, Sunderland & Sunderland, 1993: 14 (figured). Description: Shell small (10-17,7 mm), fusiform, with tall spire and moderateK elongate bod\ \\ horl gradualK tapering to short, unnotched anterior canal (w = 0,34- 0.45, a = 0,31-0.47). Whorls rounded, lacking sulcal region. Sculpture of axial ribs, 10-12 on penultimate whorl, extending suture to suture, tending to be onK slightly curved below the suture at area usually occupied by sulcus, and fine spiral striae of "butterfly wing" type overall. Sinus moderately deep, U-shaped, occupying most of shoulder slope, with parietal callus. Varix 1/4 whorl behind thin lip \\ ith shallow stromboid notch. Color white, with variable banding of orange shades. Operculum leaf- shaped, with terminal nucleus (figure 33). Type material: Holot\pe, USNM 880073. Parat\pes, one each at NMNH, AMNH, MCZ, DMNH, ANSP, LACM, \M, MORG, NHM, MNHN, IRSNB; others in author s collection. Type locality: Gulf of Mexico west of Crystal River, Florida, 30 fm (59 m), dredged, Jim Moore!, May, 1963. Range: Known onK from the Gulf of Mexico from the Cedar Keys to the Florida Ke\ s. The shell figured b\ the Sunderlands appears to be this species but would need verification as to identification and locality. A number of specimens have lost the precise locality data but are from the Gulf. All material is from the Jim Moore dredg- ings. Remarks: The author initialK' considered F, petiti a possible dwarf form of F, janetae on the basis of general appearance. However examination of additional mate- rial showed the resemblance to be superficial. The taxon is nearer a species that, on cursor\ scrutiny, does not even appear to be of the group, Fenimorea pagodula. (See below.) The major differences between the two spe- cies are that F. petiti is more tumescent, although quite variable, and has fewer, wider, and straighter axials. Fenimorea petiti shows variation of color pattern, the spiral band being pale and perhaps interrupted in some examples, but this is nearK alwaxs stronger and more colorful than the bland and unremarkable banding of F. pagodula. The species is named for Mr. Richard E. Petit for his kind donation of the material and in recognition of his many contributions to malacology. Fenimorea pagodula (Dall, 1889) new combination (Figure 3) Drillia pagodula Dall, 1889:90, pi. L3, fig. 6. Remarks: Identification of F. pagodula has been on uncertain grounds because Dall did not designate a ho- lot\ pe and a number of s\ ntype lots w ith shells of some- what disparate nature, located both at the USNM and MCZ, exist. These have been examined and the shell illustrated in figure 3 (USNM 87471, ofi^ Havana, 119- 175 fm (217-320 m)) is a good representation of Dalls description and figure despite being two mm shorter than the measurement given by Dall in the plate caption. The width of the specimen illustrated in figure 3 is 5.8 mm, as is stated in Dall s description. It is here designated lectotype of the species. .Although not bright in general appearance, as t\pical of the Fenimoreas, this species has the microsculpture characteristic of Fenimorea. Paralectotypes: USNM 887466 (separated from USNM 87471); USNM 87472, 3 specimens, Dominica, 118 fm (216 m), sand, 65°F, Blake St. 177; MCZ 7068, off Bar- bados, 154 fm (282 m), Blake St. 282; MCZ 7069, 11 specimens (a mixed lot and only 5 specimens are true F. pagodula). off Barbados, 103 fm (188 m), Blake St. 273; MCZ 7070, off Barbados, 73 fm (133 m), Blake St. 290; MCZ 7071, W. of Fla., 50 fm. Genus SedUiop.sis Petuch, 1988 Type species: Pleurotoma gracilis Conrad, 1830. SedUiopsis riosi Tippett, new species (Figures 6, 7) Description: Shell small (12-15 mm), claviform, tur- reted, with moderately tall spire, short, unnotched si- phonal canal (w = 0.37, a = 0.41). Whorls about 7 in- cluding protoconch of two whorls, the first smooth, the second bearing 3-4 spiral cords. Adult whorls rounded, somewhat tumid, with distinct suture followed by sub- sutural cord, deep shoulder sulcus, whorls angulated by upper ends of axial ribs. Sculpture of dominant axial ribs crossed b\ incised spiral striae producing w ider, some- w hat flattened cords most evident as lateralK elongate Page 134 THE NAUTILUS, Vol. 109, No. 4 beading on ribs. Hibs broad, wi-ll rounded, about 16 on bod> wliorl, with a tendency to bifurcate basally. Sinus moderately deep, U-shaped, occupying most of sulcus, apex at mid-point. No stromboid notch or varix. Color uniform mi!k\ white. Types: Holotype, MORG 31.775. Paratypes, MORG 31.776 -one specimen, MORG 31.777 - Bve specimens; USNM 880075 - one specimen; one specimen in author's collection. Type Locality: Off Sao Paulo, Brasil, 24°31'S,44°28"W, in 250 m. This species and Lioglyphostoma hendersoni, below, were dredged using a beam trawl b\- the "W. Besnard" on Dec. 9, 1988, Bottom temp. 13.2°C. Ana Maria Vaninl Range: Known by onK the type material. Remarks: This species represents the discovery of a living member of what has hitherto been known solely as a fossil genus. The distinguishing feature of the genus is the characteristic protoconch (figure 8), which bears three or four spiral cords on the second and, if present, third whorl(s), a state unique for the family in the western .\tlantic. Gibson 1,1962:238), discussing Conrad's Pleurotoma gracilis, recognized this and identified sev- eral (fossil) species with the same type of protoconch, but onl\ suggested that the group mav deserve ranking as a separate genus or subgenus. Petuch (1988) erected the genus, feeling that the closest relative was Sedilia Fargo (1953:370), a Pliocene genus from southwestern Florida having a different type of protoconch. He stated, based on the available knov\ledge, that Sediliopsis be- came extinct after the time of the Saint Mary s formation (late Miocene of the Maryland coastal plain). The present species apparently represents an instance of relict-pocket occurrence, as discussed b\ Petuch. Further study of forms with this type of protoconch shows that others outside the restricted geotemporal range of S. gracilis (middle western Atlantic coastal plain Miocene) exist:S. chowanensis (Gardner, 1948), from the upper Pliocene of North Carolina and S. aphanitoma (Dall, 1892) and S. ondulum (Fargo, 1953), both from the Pliocene of Florida. This broader geographic and temporal range of members of the genus makes it reasonable that Sediliop- sis could also occur in the Recent tauna of Brasil. This species is named in honor of Professor Eliezer de C. Rios for his contributions to Brasilian malacology and for pro- viding the material for study. Subfamily Crassispirinae Morrison, 1966, ementled Mc- Lean, I9'71 Genus Lioglyphostoma Woodring, 1928 Type species: Lioglyphostoma adcniatuiu Woodring, 1928. Lioglyphostoma hendersoni (Bartsch, 1934) (Figures 9, 26, 34) Glyphostoma (Glyphostomops) hendersoni Bartsch, 1934:17, pi. 5, figures 2,5,8. Sp."E" Rice & Kornicker, 1965:128, pi. 7. tig. 18. Material examined: See with Sediliopsis riosi above; 34 specimens. Voucher specimen, L;SNM 880078; 2 spec- imens in author's collection; remainder at MORG. Remarks: Described from Puerto Rico, this species has also been recorded from off North Carolina (Porter, 1975: 35). The present report of L. hendersoni from off Brazil and a specimen from the Campeche Bank. Yucatan (Rice & Kornicker, 1965) indicate that it is widespread. The shells were compared with both the holotype and ap- proximately 65 other lots in the USNM from the Miami and Florida Keys areas, .\lthough the present material shows stronger beading at the intersections of the axial and spiral sculpture than does the holotype, there is com- plete intergrading of forms. Bartsch (1934) selected a specimen with weaker sculpture as the type. Unknown previousK', the operculum (figure 34) is leaf shaped \\ ith a terminal nucleus, and the radula (figure 26) show s about 40 pairs of duplex teeth with a presumably detached accessor)' limb (detachment can not be determined con- clusi\eK using light microscopy ). Genus Inodrillia Bartsch, 1943 Type species: Pleurotoma (Drillia) nucleata Dall, 1881. Inodrillia nucleata (Dall, 1881) (Figure 20) Pleurotoma (Drillia) nucleata Dall, 1881:9. Drillia nucleata Dall, 1889:92, pi. 11, fig. 1, Inodrillia (Inodrillia) nucleata Bartsch. 1943:102, pi. 8, fig. 6; pi 10, fig. 4; pi 13, fig. 1. Genus Viridriltia Bartsch. 1943 Type species: Viridrillia nilliami Bartsch, 1943. Viridrillia williami Bartsch, 1943 (Figure 21) Viridrillia (Viridrillia) willianu liartsch, 1943:92. pi. 8. fig. 2; pi. 10, fig.2, Viridrillia hendersoni Bartsch, 1943 (Figure 22) Viridrillia { Viridrillina) hendersoni Bartsch, 1943:99. pi. 7. fig. 3; pL 10, fig, 1: pi, 15. fig. 5, Remarks: Bartsch's genera and the species from them are noted to ensure placement in the subfamily tirassis- pirinae and to illustrate the radulae of Viridrillia williami and V. hendersoni. Powell (1966, text figs. E 109-111) had figured J.P.E. Morrison's dray\ings of radulae, from the USNM, of Inodrillia nucleata. I. miamia. and I. ino Bartsch, but not those ot Viridrillia. although they yvere D L, Tippett, 1995 Page 135 available to him. Powell commented about Viridrillia (1966:74, with the genus Cerodrillia): "... a very dif- ferent raduia. which consists oni\ of a pair of elongate, fohated marginals.' He placed the genera in his subfam- il\ Cla\ inae, as he interpreted it at tliat time, principalK on the basis of shell characters. SubsecjuentK, McLean (1971:119) separated this grouping, largeK on the basis of radular form, into the subfamilies Ciavinae [subse- quentk DriUiinae, Claxinae being preoccupied (Cer- nohorsks , 1985:60)] and Crassispirinae. The teeth of Inodrillia and Viridrillia, although not typical, are most similar to those of Crassispirinae, certainly not to those of the Driiliinae. Sysoev. in Taylor et al. (1993:164), recognized this and listed Inodrillia as a crassispirine genus. Morrison's illustrations are somewhat unclear so I reviewed his slides and the figures presented here were made. The teeth of both genera are essentiali\ the same, consisting of simple, elongate, solid, pointed marginals. They are not clearK duplex but show a lengtliw ise thick- ening suggestive of an accessory limb, perhaps a step in the evolution towards accessor) limbs. Bartsch's subgen- era Viridrillina, Inodrillina. Inodrillara now are s\n- onymized with their respective parent genera Viridrillia and Inodrillia. Subfamily Clathurellinae H. & A. Adams, 1858 Genus Clathurella Carpenter, 1857 Type species: Clavatula rava Hinds, 1843, by subse- quent designation Cossmann, 1896. Clathurella eversoni Tippett, new species. (Figures 10, 11) Description: Shell \ery small (4.5 mm), biconic-fusi- form, with tall spire, short anterior canal (w = 0.38, a = 0.33). Protoconch of three smooth whorls. Adult whorls four, sculptured by 11-12 rounded axial ribs separated by equal interspaces, crossed by spiral cords forming laterally elongate beading at intersections. Spiral cords extend down over base and canal. Sinus moderately deep, U-shaped, on shoulder slope, with onh' modest parietal callus. Enlarged axial rib behind lip. Five spiralK' elon- gate denticles inside outer lip. Columella with two weak pustules. Color light brow n w ith paler band on peripheral spiral. Type: Holot) pe, USNM 880074. Type locality: Off Dania Beach, Florida, between sec- ond and third reefs. 70 ft (21 m), in rubble, SCUBA at night. Gene Eversoni Remarks: .although know n only from the holotype, this species is distinctive and warrants description. The ho- lotype is still slightK juvenile and would undoubtedly show a deeper sinus at full maturity; also it is somewhat worn. There is a vestige of carination on the later protoconch whorls and traces of minute granulation on the shell surface, which are characteristic of the genus. L ntil the description of C. fuscoba.iis Rehder (1980:87, pi, 11, figs. 3,4) from Easter Island, Clathurella in its modern concept was known only from the tropical east- ern Pacific. The present species extends the range of the genus to the western Atlantic area. Clathurella eversoni is nearest C, rava (Hinds, 1843), differing in being small- er, having a broader shoulder slope, stronger spiral cord at the shoulder, and different color pattern. This species is named for its discoverer, Mr, Gene Everson. Subfamily Zonulispiriiiae McLean, 1971 Genus Pilsbryspira Bartsch, 1950 Type species: Pilibrijfipira pilsbryi Bartsch, 1950 ( = Plcurotoma jaijana C, B, Adams, 1850b). Pilsbryspira (Pilsbryspira) albocincta (C. B. Adams, 1845) (Figures 27,29) Pleurotoma albocincta C B, .\dams, 1845:3 (reprinted Clench & Turner, 1950:25:3, pi. 29, fig, 5), Material exainined: Two specimens, alcohol pre- served. Cabbage Patch, Abaco Island, Bahamas, under rocks in 3 ft water, Colin Redfern!, July 8, 1972. Remarks: Shells are an exact match to the lectotype (Clench & Turner, 1950, pi. 29, fig. 5). Animal is uni- formly grayish pink. Prominent inhalant siphon leads to a large gill. Proboscis is of the intraembolic form. A much convoluted poison gland w ith a large muscular bulb is present. The oesophagus was empt\ . Raduia (figure 27) consists of numerous pairs of broad marginal teeth, flar- ing at the distal end and bearing a small barb. The teeth are t\pical of the subiamiK'. Subgenus Pdsbryspira (Syniphispira) McLean. 1971 Type species: Crassispira nymphia Pilsbry & Lowe, 1932. Pilsbryspira (Nymphispira) aubcrti (Lam\', 1934), new combination (Figure 19) Drillia auberti Lamy, 1934:435 (description and figure), Crassispira auberti (Lamy, 1934) Radwin, 1969:2:32 (list), Pilsbri/spira harjordiana (Reeve, 1843) N'okes & N'okes, 1983: ;30, pi, 2, fig, 11a, Crassispira harjordiana (Reeve, 1843) Sunderland & Sunder- land, 1993:14 (figure). Material examined: 3 specimens, dredged, 120 ft (36 m), Porto Bello Ba\ , Caribbean Panama, James Ernest! N'oucher specimen, USNM 880079. Others in Everson and Ernest collections. Remarks: .As seen in the synonymy, there has been uncertainty as to the identification and placement of this species. Lamy's type material cannot be found (pers. comm,, P. Bouchet) and thus identification must neces- Page 136 THE NAUTILUS, Vol. 109, No. 4 saril) be based on comparison with Lam\ s description and figure. Fortunately, these are reasonably good, and the present material appears to be this species, agreeing in overall form, sculptural detail, and color. Also the current material is from the same geographic area. The material referred to in the cited publications appear to be the same species, it can be assumed that Radwin U969) was accurate in his identification because his ma- terial came from the t\ pe localitv . A broader geographic range than known previousK is suggested. The N'okes' (1983) specimens came from Yucatan, not a surprising range extension, while Sunderland and Sunderland (1993) reported on material from Cuba. Identification of the Cuban specimens as Crassispira (Glossispira) harfordi- ana is in error, despite superficial similarity. This species is from the eastern Pacific and has a more tumescent outline, crassispirine sinus structure, blackish color, and other, finer, differences. Although soft parts of P. auberii are not available, the sinus structure, beaded base, and color pattern are consistent \\ ith the present assignment. Subfamily Mangeliinae P. Fischer, 1887 Genus Pyrgocythara Woodring, 1928 Type species: Pyrgocythara eminula Woodring, 1928. Pyrgocythara filosa Rehder, 1943 (Figure 25) Pyrgocythara filosa Rehder, 1943:203, pi. 20, fig. 2. Material examined: Six specimens, alcohol preserved. Pine Island, east Florida, sieved from Spartina grass, 24 September 1993, John Wise! Remarks: Shells of these specimens match the holotype except that the axials are slightK' narrower and their shoulder angulation is slightly sharper. Animal white with black eyes located near the ends of short stalks. There is a prominent inhalant siphon. The mantle is speckled black and this coloring extends back over the visceral segment. There is a coiled poison gland and muscular bulb. The radula consists of paired marginal teeth enclosed in a radular sac. Radular tooth (figure 25) elongate, sharply pointed, bearing a small barb 1/3 the distance from the tip, and has a projection at the base from which a triangular extension of tissue protrudes posteriorly. The radular teeth are of the "hilted-dagger ' type described by Powell (1966) and are very similar to those of Pyrgocythara danae (Dall. 1919) from the east- ern Pacific except that the teeth of that species do not have barbs (figure 24). Pyrgocythara plicosa (C. B. Adams, 1850) (Figure 23) Pleurotoma plicosa C B. Adams, 1850b:54 (new name for Pleurotorna plicata C. B. .\dams, 1840, non Lamarck, 1804). Material examined: 2 live specimens, Little Jim Island, Fort Pierce, Florida, grass and mud flats, intertidal, hand sieved, 24 la 25 Januar\ 1991, John Wise! Remarks: The animal has a whitish, semitranslucent foot and head speckled with fine, bright, ivor\-white spots, the eyes prominent toward the ends of rather long stalks. No operculum. The mantle is w hite speckled w ith sooty-black, and this coloration extends back over the visceral lobe. .A long penis with a fingernail-shaped end is attached to the right side of the head and curls back inside the mantle cavity. Large gills, osphradium, and hypobranchial glands are present. The proboscis is in- traembolic. The radular teeth i figure 23) are enclosed in a radular sac and immber about 35 pairs. They are of the "hilted-dagger" form and resemble those of P. danae but lack an angulation toward the tip. ACKNOWLEDGEMENTS The author thanks the USNM and the staft for the op- portunity of working with the collection and use of equipment and facilities. Appreciation is extended to Dr. John Wise for supplying material, and to him and Dr. Jose Leal for assistance and support during the work. I also thank Dr. Alan Kabat for suppKing information concerning Dall s "Blake" material and kindK review ing this paper. Professor Eliezer de C. Rios of the MORG supplied material of Sediliopsis riosi and Lio- glyphostoma hendersoni. Dr. J. \an Goethem of the IRSNB loaned the lectot\ pe of Drillia dautzcnbergi. Dr. Douglas Wolfe contributed specimens of Drillia wolfei. Mr. Richard E. Petit supplied a number of specimens of Fcnimorea petiti. Mr. Gene Everson and Mr. Colin Red- fern supplied specimens of Clathurclla cveraoni and Pils- bryspira albocincta respectively. Mr. E\erson supplied the voucher specimen of Pilsbryspira auberti. Tan\a Kausch of the MCZ supplied t\pe material of Drillia pagodula. The author thanks these indi\ iduals. Opercula and final radula and animal illustrations were draw n by Mathilde Duftv. LITERATURE CITED .Abbott, R, T 1974. .\merican Seashells, 2nd Ed. New York, etc. \'an N'ostraiid Reinhold Co. 663 pp. Illustrated. Adams, C. B. 1840. Descriptions of thirteen new species of New England shells. Boston Journal of Natural History 3;318-332, Reprinted, C;leiich & Turner, 1950. Adams, C. B. 1845. Specierum novarum conchyliorum, in Jamaica repertorum, synopsis. Proceedings of the Boston Society of Natural History 2:1-17 Reprinted, Clench & Turner, 1950. Adams, C. B, 1850a- Notes on tlies\non\ m\ ol certain marine shells Contributions to Conchology no. 4:54-55 Reprint- ed, Clench in Turner, 1950. Adams, C. B. 1850b. Descriptions of supposed new species D. L. Tippett, 1995 Page 137 of marine shells which inhabit Jamaica. Contributions to Conchologv no. -4:56-68 Reprinted, Clench & Turner, 1950. Adams, H. & .\. .•\danis. 1858. The genera of recent mollusca. 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Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U. S. Coast Survey Steamer "Blake, Lieut-Commander C. D. Sigsbee, U. S, N., and Commander J. R. Bartlett, U. S. N., commanding. 29, -Report on the Mollusca. Pt. 2. Gastropoda and Scaphopoda Bulletin of the Museum of Comparative Zoologv, at Har\ard College 18:1- 492, pis. 10-40. Dall, W. H. 1892. Contributions to the Tertiary fauna of Florida, with especial reference to the Miocene Silex-beds of Tampa and the Pliocene beds of the Caloosahatchie River. Transactions of the Wagner Free Institute of Sci- ence of Philadelphia. 3(2):201-44S, pis. 13-22. Dall. W. H. 1918. Notes on the nomenclature of the mollusks of the family Turritidae. Proceedings of the United States National Museum 54(2238):313-333. Dall, W. H. 1919. Descriptions of new species of mollusks of the famil> Turritidae from the west coast of America and adjacent regions. Proceedings of the United States National Museum 56(2288):l-86, pis. 1-24. Dautzenberg. P. 1900. Croisieres du yacht Chazalie dans I'At- lantic. Mollusques. Memoires de la Societe zoologique de France 13:145-265 (1-121), pis. 9,10. Fargo, W G. 1953. The Pliocene Turridae of Saint Peters- burg, Florida. In: Olsson, A A. & .A. Harbison, Pliocene Mollusca of Southern Florida, The Academy of Natural Sciences of Philadelphia Monograph 8(2):363-409, pis. 1-24. Fischer, P. 1887. Manuel de Conchyliologie et de Paleon- tologie Conchyliologique ou histoire naturelle des moll- usques vivants et fossiles .... Paris. F. Savy. 1369 pp., 23pls., 1138 text figs. Gardner. J. 1948. Mollusca from the Miocene and Lower Pliocene of Virginia and North Carolina Pt. 2. Scaphopoda and Gastropoda. United States Geological Survey Profes- sional Paper 199-B:79-310, pis. 24-38. Gibson, T. G. 1962. Revision of the Turridae of the Miocene St. Mary s formation of Maryland. 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December 1986; Part 4. Subfamilies Drilliinae, Crassispirinae and Strictispi- rinae, 29(1): 167-320, 284 figures. May 1988; Part 5. Sub- family Taraninae, 32:325-339, 21 figures, Octoberl991; Part 6. SubfamU) Mangeliinae, section 1, 33(2):461-575, 193 figures, October 1992; Part 6. Subfamily Mangeliinae, section 2, 34(2):317-367, 88 figures, October 1993; Part 7. Subfamily Crassispirinae, section 2, 35: 177-228, 95 figures, October 1994 (article title now headed "Turridae [s.l.]" subsequent to proposed classification of the conventional family Turridae b> Taylor et al., 1993). Kohn, A. J. and J. H McLean. 1994. Review of "'Foregut anatomy, mechanisms, relationships and classification of the Conoidea (=Toxoglossa) (Gastropoda) " bv Taylor, J. D., Y. I.Kantor & A. V. Svsoev. 1993. The Veliger 37(4): 432-435, Lamarck, J. B. 1804. Memoires sur les fossiles des environs de Paris Genre 25,Pleurotoma. Annales du Museum National d'Histoire Naturelle 3:79-90. Reprinted by Pa- leontological Research Institution, 1978. Lamy, E. 1934. Coquilles marines recueillies par M. E. Aubert de la Riie dans 1 Amerique du Sud. Bulletin du Museum d'Histoire Naturelle, Paris, ser. 2, 6(5):432-435, 1 figure. McLean, J. H. 1971. A revised classification of the family Turridae, with the proposal of new subfamilies, genera, and subgenera from the eastern Pacific. The Veliger 14(1): 114-130, 142 figs. 1 table. Maury, C. J. 1917. Santo Domingo type sections and fossils, Pt. 1. Bulletins of American Paleontology 5(29):1-251, pis, 3-39. Reprinted Kraus Reprint Corporation, 1964. Page 138 THE NAUTILUS, Vol. 109, No. 4 Morrison. J. P. E. 1966. On the families of Turridae. American Malacological Union, Bulletin for 196.5, 32:1-2. Olsson, A. .\. 1964. Neogene Mollusks from Northwestern Ecuador. Ithaca, N.Y. Paleontological Research Institu- tion. 2.56 pp., 36 pis. Petuch, E. J, 1987. 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