ATL, abit i i ark eet We in oa3 Ca ies a. prety penocers aay Mea gm Ty eG tata aL LE. ae ELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler, Founding Editor Volume 43 ney O Vie IGG NE Sao aes, ‘ ISSN 0042-3211 January 3, 2000 CONTENTS The anatomy and systematics of Latiromitra, a genus of tropical deep-water Ptychatractinae (Gastropoda: Turbinellidae) RE TPIBEE GD OU CEE ANDI GURT EMCANGT ORS sia jatorcriuctlel ley alete sista ate) opscaepe Mas levieis, sale ola: sha Sex change, reproduction, and development of Crepidula adunca and Crepidula lingulata (Gastropoda: Calyptraeidae) IRACISRIE, CROTON ys, 1S See eshte pia oe mnie Nr RI USEP OO nua UPDE rel LU ecru ST On Shell growth of Mytilus trossulus Gould, 1850, in Port Valdez, Alaska ARN VED EAN CHARD AND Ed OWARD I My FEDER (oye hele cieiais tic ile !ajie spc) e cee seep 6 @ a .0 38 Two new Neogene species and the evolution of labral teeth in Concholepas Lamarck, 1801 (Neogastropoda: Muricoidea) SIDET@ MASS Bal ENSRIP Seana ieer ey ae Meet ecu nla ets COW etal conden eae MEAS ta alta gin Pisidium tavaguyense and Pisidium pipoense, new species from northeastern Argentina (Bivalvia: Sphaeriidae) ERISIGUAN Ee SWART E eibr pre ben ar anes oats tah it, Mallar Aik ies ee aden eh Aide ak Development and metamorphosis of the planktotrophic larvae of the moon snail, Polinices lewisii (Gould, 1847) (Caenogastropoda: Naticoidea) ROBERTA AKeaEDERSENGAND EOUISESRes PAGE Mine evan areata ances Sheainte aiers iefeet allele Polygyrid land snails, Vespericola (Gastropoda: Pulmonata), 3. Three new species from north- ern California BARRVGR OM EWAN DA AICDERGD pV ITEGER Wiis seya\ls holeuel. toleiley ate lleva i=) eye) cto laiie) «a ayele!'-/ al ears CONTENTS — Continued 24 34 43 Sill 58 The Veliger (ISSN 0042-3211) is published quarterly in January, April, July, and October by the California Malacozoological Society, Inc., % Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. Periodicals postage paid at Berkeley, CA and additional mailing offices. POSTMASTER: Send address changes to The Veliger, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. Number 1 THE VELIGER Scope of the journal The Veliger is an international, peer-reviewed scientific quarterly published by the Cali- fornia Malacozoological Society, a non-profit educational organization. The Veliger is open to original papers pertaining to any problem connected with mollusks. 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Send manuscripts, proofs, books for review, and correspondence regarding editorial matters to: Dr. Barry Roth, Editor, 745 Cole Street, San Francisco, CA 94117, USA. ® This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). THE VELIGER © CMS, Inc., 2000 The Veliger 43(1):1—23 (January 3, 2000) The Anatomy and Systematics of Latiromitra, a Genus of Tropical Deep-Water Ptychatractinae (Gastropoda: Turbinellidae) PHILIPPE BOUCHET Muséum national d’ Histoire naturelle, 55, Rue Buffon, 75005 Paris, France AND YURI I. KANTOR A. N. Severtzov Institute of Problems of Evolution of Russian Academy of Sciences, Leninski prosp. 33, Moscow 117071, Russia Abstract. The anatomy of Latiromitra Locard, 1897, is very similar to that of other representatives of the Ptycha- tractinae, notably in the short or very short proboscis, the presence of an accessory salivary gland, the ventral odon- tophoral retractor passing through the nerve ring, and the position of the buccal mass at the proboscis base in contracted position. Latiromitra differs from Ceratoxancus by its fused salivary glands (clearly separate in Ceratoxancus). Based on anatomical and conchological characters, Cyomesus Quinn, 1981, and Okinawavoluta Noda, 1980, are confirmed and/ or placed in the synonymy of Latiromitra. The genus currently comprises 10 Recent and Neogene species, three in the Atlantic, and seven in the Indo-West Pacific, all in deep water at low latitudes. Teramachia chaunax Bayer, 1971, is placed in the synonymy of Latiromitra cryptodon (P. Fischer, 1882), and the Recent Benthovoluta sakashitai Habe, 1976, is placed in the synonymy of the Pliocene Latiromitra okinavensis (MacNeil, 1961). Volutomitra? vitilevensis Ladd, 1982 is placed in Latiromitra. Three new species are described: Latiromitra paiciorum sp. nov. (New Caledonia, 960— 1100 m), L. cacozeliana sp. nov. (Vanuatu, 536-775 m), and L. crosnieri sp. nov. (Madagascar and NE of Fiji, 600— 800 m). In addition, Mitra styliola Dall, 1927, from off Georgia, USA, is tentatively referred to Latiromitra. INTRODUCTION Dall, 1890, see Bouchet & Poppe, 1995]. Bouchet & War- én (1985) examined a specimen from the Philippines and presented a drawing of the radula. Based on concholog- ical similarity with Latiromitra cryptodon, they placed Prodallia barthelowi in Latiromitra and confirmed its as- signment in the Turbinellidae. Material of the genus has since been collected alive, including specimens of the type species. In the present paper, we report on the anatomy of Latiromitra and con- firm its placement in the family Turbinellidae. We also review the species included in Latiromitra, describe new species from off Madagascar, New Caledonia, and Va- nuatu, and confirm the synonymization of Cyomesus. The present paper follows our earlier revision of Ceratoxan- The genus Latiromitra Locard, 1897, erected for the sin- gle species L. specialis Locard, 1897, was originally placed in the family Pisaniidae (= Buccinidae). Thiele (1929) transferred the genus to Vexillinae [= Costellari- idae], a position accepted by Ponder (1968), while Cer- nohorsky (1970) considered it a subgenus of Volutomitra H. & A. Adams, 1853 (Volutomitridae). Bouchet & War- én (1985) assigned Latiromitra to the Turbinellidae, based on the radula of L. barthelowi (Bartsch, 1942), and synonymized Cyomesus Quinn, 1981. Harasewych (1987), while classifying Latiromitra and Ceratoxancus Kuroda, 1952, under Turbinellidae Ptychatractinae, ex- pressed doubts about their taxonomic position, due to the lack of knowledge of the radula of their type species. cus (Kantor & Bouchet, 1997). Harasewych also disagreed with Bouchet & Warén’s Abbreviations and text conventions: ag, anal gland; (1985) synonymizing Latiromitra and Cyomesus. Thus asg, accessory salivary gland; ca, buccal cavity; ct, cte- the position of Latiromitra has remained controversial, nidium; cu, cuticle; dasg, duct of accessory salivary mainly due to the lack of anatomical data. gland; dd, dead collected specimen; ddg, duct of digestive Although known for a long period only from the ho- gland; dg, digestive gland; gL, gland of Leiblein; lv, live lotype, Prodallia barthelowi has been referred to many collected specimen; moe, glandular mid-esophagus; ne, times in the literature, and its taxonomic position changed nephridium; nr, nerve ring; od, odontophore; oe, esoph- several times. Bayer (1971) was the first to classify it as agus; op, operculum; os, osphradium; p, penis; poe, pos- a turbinellid, although he mistakenly placed it in the vol- terior esophagus; pr, proboscis; prs, rhynchodaeum (= utid genus Teramachia Kuroda, 1931 [= Calliotectum proboscis sheath); r, radular membrane; rs, radular sac; s, Page 2 siphon; sd, salivary duct; sem.gr, open seminal groove; sem.p, seminal papilla; sg, salivary gland; st, stomach; t, head tentacle; tes, testis; tu, buccal tube; vL, valve of Leiblein; vpr, ventral proboscis retractor. Repositories: BMNH, The Natural History Museum, London; MNHN, Muséum national d’Histoire naturelle, Paris; MOM, Musée Océanographique de Monaco; NSMT, National Science Museum, Tokyo; UMML, In- vertebrate Museum, Rosenstiel School of Marine & At- mospheric Sciences, University of Miami; USNM, Na- tional Museum of Natural History, Smithsonian Institu- tion, Washington DC. ANATOMY Alcohol preserved material has been obtained for two species, the anatomy of which is very similar. Specimens with dry soft parts have permitted examination of the radula in two additional species. Latiromitra cryptodon The dissected specimen (Brazil, MD55, sta. CB79) has a shell height of 21.6 mm, last teleoconch whorl height 14.8 mm, aperture height 7.7 mm, siphonal canal height 2.7 mm, shell diameter 8.4 mm. After removal of the odontophore with radula, the organs of the body hae- mocoel were serially sectioned. External anatomy (Figure 1A, B): The body consists of three whorls; the mantle cavity spans 0.75 whorl; the ne- phridium 0.3 whorl; and the digestive gland 1.25 whorl. The body is pale yellowish, it lacks pigmentation except for a small portion of the mantle near the siphon and head tentacles, which bears very small black spots. The oper- culum is large (Figure 1A—op), occupying 0.75 aper- ture height, narrow, subrectangular, very thin, flexible, transparent, and yellow. The columellar muscle is at- tached on the upper fourth of the operculum. The foot is short (L/W ~ 1.5). The siphon is short, simple, slightly protruding beyond the mantle edge. The columellar mus- cle is thick, consisting of 1.5 whorls, with three rather deep grooves, corresponding to the columellar teeth. The mantle is thin, the mantle cavity organs clearly visible through it. The mantle is short, not covering the head base and penis. The head is broad, with very short, stout ten- tacles and very large eyes. Mantle cavity: The entire mantle cavity was filled with a thick unremovable layer of hypobranchial gland secre- tion; it was impossible to examine the morphology of the mantle cavity in detail. The ctenidium is very long and occupies nearly the entire mantle cavity length, narrow (L/W ~ 7) with high hanging leaflets. The osphradium is large, 0.75 as long as the ctenidium. The anal gland is small, visible through the mantle as a narrow dark strip. Digestive system: The organs of the body haemocoel are The Veliger, Vol. 43, No. 1 compact, and their in situ position resembles that of Cer- atoxancus teramachii (Kantor & Bouchet, 1997). The contracted proboscis (Figure 1D—pr) is short (about 1 mm long, or 13% of aperture height), smooth, and oc- cupies nearly the whole rhynchodeal cavity. The rhyn- chodaeum (= proboscis sheath) is thin-walled and lined with a tall epithelium. The moderate-sized paired mus- cles, probably functioning as proboscis retractors, are at- tached latero-ventrally to the anteriormost part of the rhynchodaeum (Figure 1F—vpr). This indicates that the whole rhynchodaeum takes part in the proboscis evertion, and when the proboscis is completely everted, these re- tractors are attached to the inner wall of the proboscis. The buccal cavity is broad and lined with a thick cu- ticle. The buccal mass is large, broad, and muscular, and projects ventrally beyond the rear of the retracted pro- boscis (Figure 1F—od). The ventral odontophoral retrac- tor passes through the nerve ring, follows the bottom of the cephalic haemocoels, and joins the columellar muscle. The radula (Figure 3A—C) is about 1.6 mm long (7.4% of shell height and 20% of aperture height) and about 107 wm wide (0.49% of shell height and 1.39% of ap- erture height). The radula projects beyond the rear of the proboscis, and consists of more than 80 transverse rows. The rachidian teeth (Figure 3B) bear three cusps that are curved in profile, the central one 1.5 as long as the lateral cusps. The cusps emanate from close to the anterior edge of the basal part. The basal part of the rachidian has a rather deep notch on the anterior edge. The lateral teeth are unicuspid, with a long narrow base. The length of the lateral tooth base equals 0.80 the rachidian width. After leaving the proboscis the esophagus forms a very short loop before opening into the valve of Leiblein (Fig- ure 1D—vL). The valve of Leiblein is well defined, much broader than the esophagus, and oval-pyriform. It has a conical ciliar valve, similar to that in Muricidae. Between the valve and the opening of the gland of Leiblein the esophagus is widened and has well-developed dorsal glandular folds. This part, representing the mid-esopha- gus, is long (ca. 2.3X proboscis length). The posterior esophagus runs along the left side of the gland of Leiblein and opens into the stomach. The stomach is very small by comparison to the an- terior foregut, and broadly U-shaped. The fixation of the specimen does not permit examination of the inner anat- omy of the stomach. Judging from the external view, the stomach has a small caecum. Typhlosoles of even size can be seen through the stomach wall. The gland of Leiblein is large, gray in the fixed spec- imen, tubular, coiled anteriorly, and simple posteriorly. The gland opens into the esophagus through a duct with- out defined constriction (Figure 1E). The salivary glands are fused, compact, flattened, and cover the posterior proboscis sheath dorsally. When the organs of the body haemocoel are extended, the glands P. Bouchet & Y. I. Kantor, 2000 Page 3 Figure 1 Anatomy of Latiromitra cryptodon, male (Brazil, MD55, sta. CB79, shell height 21.6 mm). A, B. Body, removed from the shell. C. Anterior part of the body with the mantle removed to show the penis and seminal groove. D. Organs of the body haemocoel, expanded. E. Opening of the gland of Leiblein into the esophagus, from the ventral side. E Proboscis sheath with odontophore, from the ventral side. C—F in the same scale. Page 4 The Veliger, Vol. 43, No. 1 lie above the nerve ring and embrace the valve of Lei- blein (Figure 1D—sg). Immediately after leaving the glands, the salivary ducts enter the walls of the esophagus in front of the valve of Leiblein. The ducts follow in the walls of the esophagus to their opening in the buccal cav- ity (on the drawing they are shown by dashed lines). There is a medium-sized unpaired accessory salivary gland, situated on the right side of the nerve ring (Figure 1D—asg). Gland histology is typical for neogastropods, consisting of a thick outer epithelial layer and a thin inner one, delimited by a layer of circular muscle fibers. The duct of the gland opens ventrally in the anterior part of the buccal tube. The highly concentrated circumesophageal nerve ring is situated in the position typical for Muricoidea, just pos- terior to the valve of Leiblein (Figure 1D—nr). Male reproductive system (Figure 1C): The dissected specimen was a mature male. The pallial portion of the system is represented by a muscular open groove, running anteriorly to the base of the rather long penis and then along its inner lateral edge to a medium-sized, conical papilla. Latiromitra barthelowi The largest specimen was anatomically studied (Indo- nesia, KARUBAR, sta. CC41). It has a shell height of 52.0 mm, last teleoconch whorl height 31.3 mm, aperture height 17.4 mm, siphonal canal height 6.0 mm, shell di- ameter 17.7 mm. The foot with the anterior foregut was serially sectioned. The radula has been studied on a spec- imen from Indonesia (KARUBAR, sta. CP72). It has a shell height of 48.7 mm, last teleoconch whorl height 29.0 mm, aperture height 16.4 mm, siphonal canal height 5.5 mm, shell diameter 15.3 mm. External anatomy: The upper part of the body was torn off during extraction from the shell. The remaining part consists of 1.3 whorls; the mantle cavity spans 0.75 whorl. The body is pale yellowish, lacking differential pigmentation. The operculum is small, about 0.3 aper- ture height, narrow-leaf shaped and slightly curved, with terminal nucleus (Figure 2B). The foot is short (L/W ~ 1.5). The siphon is short, simple, slightly protruding be- yond the mantle edge. The columellar muscle is thick, consisting of about 1.5 whorls, with two deep grooves, corresponding to the columellar teeth. The mantle is rath- er thin, covering the head base, the mantle cavity organs clearly visible through it. The head is broad, with long cylindrical tentacles and large eyes. Mantle cavity: The ctenidium is very long and occupies nearly the whole mantle cavity length, narrow (L/W ~ 10.5), with high hanging leaflets. The ctenidial lamellae have the form of tall triangles. The osphradium is large, 0.5 as long and 1.5 as wide as the ctenidium, asym- metrical with the right side nearly twice as broad as the left. The hypobranchial gland is covered with a thick mu- cus layer and is transversely plated. The rectum opens close to the mantle edge. The anal gland is not visible. Digestive system: The general arrangement of the ante- rior digestive system is very similar to that of Latiromitra cryptodon. The proboscis is short, about 2 mm long (11% of aperture height), with folded walls, and occupies about two-thirds of the rhynchodeal cavity. The rhynchodaeum is thin-walled, and lined with very tall epithelium. The moderate-sized paired muscles, func- tioning as proboscis retractors, are attached ventrally to the anterior part of the rhynchodaeum and join the colu- mellar muscle. The buccal mass is large (Figure 2A—od), broad, and muscular, projecting ventrally beyond the rear of the re- tracted proboscis. The radular diverticulum opens in the posterior part of the proboscis into the broad buccal cav- ity. The buccal tube (Figure 2A—tu) is lined with very thick cuticle (Figure 2A—cu), and leads from the mouth opening to the buccal cavity (Figure 2A—ca). The length of the buccal tube is at least one-third of the length of the proboscis. The radula (Figure 3D—F) is about 4.6 mm long (9% of shell height and 28% of aperture height), about 250 wm wide (width 0.51% of shell height and 1.4% of ap- erture height) and consists of about 130 rows. The rach- idian tooth, with a short arched base (Figure 3E), bears three short obtuse cusps, emanating from close to the an- terior edge of the basal part, the central cusp being lon- gest and widest. The rachidian basal parts have a shal- lowly arched anterior edge and a rounded posterior edge, which is overlapped by the following tooth. The posterior corners of the rachidian tooth are narrow, thin projections (Figure 3E, F) that join the subradular membrane. The laterals are similar in shape to those in L. cryptodon. The length of the lateral tooth base equals 0.60 the rachidian width. The subradular cartilages are paired, not fused anteri- orly, projecting beyond the base of the retracted probos- cis. The thin ventral odontophore retractor passes through the nerve ring to join the columellar muscle. After leaving the proboscis, the esophagus forms a very short loop before opening into the large oval valve of Leiblein. The dorsal glandular folds of the mid-esoph- agus are well developed. The gland of Leiblein is large, tubular, coiled in its anterior part, and simple posteriorly, opening into the esophagus without visible narrowing of the duct. The inner cavity of the gland of Leiblein is subdivided by the projections of the walls into numerous small chambers. The salivary glands are fused, large, and acinous. The border between them is not visible. They are compact, flattened, situated above the nerve ring, and embrace the valve of Leiblein, also covering the posterior proboscis sheath dorsally. Immediately after leaving the glands, the P. Bouchet & Y. I. Kantor, 2000 sie 5 cS Ry; Z cunnmuiYZ (a VW Y, Page 5 prs Figure 2 A. Longitudinal semidiagrammatic section of the proboscis of Latiromitra barthelowi (Indonesia, KARUBAR, sta. CC41, male, shell height 52.0 mm), scale bar 0.5 mm. B. Operculum from the inner side (at the left) and from the outer side (at the right). The place of columellar muscle attachment is dotted, scale bar 2 mm. C. Stomach from the outside; the dashed line represents the border of the lobe of the digestive gland; the openings of the ducts of the digestive gland at the inner surface of the stomach are shown by dotted circles, scale bar 2 mm (B, C, Indonesia, KARUBAR, sta. CP39, shell height 47.2 mm). D. Operculum of a paratype of L. crosnieri Bouchet & Kantor, sp. nov. from the outer side (at the left) and from the inner side (at the right). The place of columellar muscle attachment is dotted, scale bar 2 mm. salivary ducts enter the walls of the esophagus in front of the valve of Leiblein. The ducts follow in the walls of the esophagus to their opening in the buccal cavity. The place of opening of the ducts is shifted anteriorly from the opening of the radular diverticulum. One duct opens ventrally, the other dorsally. Near their opening, the ducts are greatly widened and form the ‘‘ampullas”’ lined with smooth epithelium (marked by the arrows on Figure 2A). The Veliger, Vol. 43, No. 1 Figure 3 Radulae of Latiromitra spp. Latiromitra cryptodon-A—C) (Brazil, MD55, sta. CB79): A. Dorsal view of the radular ribbon. B. Enlarged rachidian teeth. C. Bending plane of the radular ribbon. Latiromitra barthelowi (D—F) (Indo- nesia, KARUBAR, sta. CC72): D. Dorsal view of the radular ribbon. E. Enlarged rachidian teeth. F Enlarged posterior corners of the rachidian tooth. Arrows on E, F show projections that join the subradular membrane. Scale bars 100 wm (D); 50 pm (A, C, E); 20 wm (B); 10 pm (F). These ampullas open into the buccal cavity with rather ventral sides of the proboscis sheath. The gland histology narrow ducts. is typical for the neogastropods, consisting of a thick out- The accessory salivary gland is very large, tubular, er epithelial layer and a thin inner one, delimited by a coiled anteriorly, and situated at the right-posterior and layer of circular muscle fibers. The duct of the gland P. Bouchet & Y. I. Kantor, 2000 Page 7 opens ventrally in the anterior part of the buccal tube (Figure 2A—dasg). The stomach (KARUBAR, sta. CP39) is relatively small, broadly U-shaped with a short caecum (Figure 2C). The posterior part of the esophagus and its entrance into the stomach is covered with the lobe of the digestive sys- tem (the border of the lobe is marked by dashed line on Figure 2C). The ducts of the digestive gland are paired and small. The first is situated near the entrance of the esophagus into the stomach, the second near the opening of the rectum (shown by dashed circles in Figure 2C). Male reproductive system: One of the specimens with a dried body (KARUBAR, sta. CP39) was rehydrated and appeared to be a mature male. The pallial portion of the system is represented by a muscular open groove, running anteriorly to the base of the penis (which is as long as the entire mantle cavity) and then along its inner lateral edge to the tip. The poorly preserved material does not show details of the penis tip, but it seems to be similar to that of Benthovoluta claydoni (Harasewych, 1987: fig. 26). Latiromitra crosnieri sp. nov. The operculum of the paratype (shell height 30.4, ap- erture height 12.4 mm) is about 3.6 mm (0.30 aperture height), transparent, yellowish, narrowly leaf-shaped, and with a terminal nucleus, which is missing (Figure 2D). The radula of one paratype was studied (Figure 4A— C). It is about 2.7 mm long (9% of shell height and 17% of aperture height), and consists of about 105 rows, nar- row, width about 115 wm (0.38% of shell height and 0.92% of aperture height). The rachidian teeth (Figure 4B) bear three cusps that are curved in profile, the central one 1.3X as long as the lateral cusps. The cusps emanate from close to the anterior edge of the basal part. The basal parts of the rachidian have a shallow notch on the anterior edge and a rounded posterior edge, which is overlapped by the following tooth (Figure 4B, C). The laterals are similar in shape to those in L. cryptodon, unicuspid, with a long narrow base. The length of the lateral tooth base equals 0.70X the rachidian width. Teeth on the working surface are rather worn. On Figure 4C the teeth from the bending plane are illustrated to show the shape of the posterior edge of the rachidians. Latiromitra okinavensis The radula of the specimen with shell height 47.5 mm and aperture height 17.7 mm was studied (Figure 4D-F). It is about 3.1 mm long (6.5% of shell height and 17.5% of aperture height), and consists of about 110 rows, width about 240 pm (0.51% of shell height and 1.35% of ap- erture height). The rachidian tooth bears three short ob- tuse cusps, emanating from close to the anterior edge of the basal part, the central cusp being longest. The rachi- dian basal parts have a shallowly arched anterior edge and a rounded posterior edge, which is overlapped by the following tooth. A damaged tooth is illustrated (Figure 4F) to show the shape of the posterior edge. Laterals are similar in shape to those in L. cryptodon. The height of the lateral tooth base equals 0.70 the rachidian width. SYSTEMATICS Genus Latiromitra Locard, 1897 Type species: (by monotypy) Latiromitra specialis Lo- card, 1897 [= Mitra cryptodon P. Fischer, 1882], Recent, tropical Atlantic, bathyal. Synonyms: Okinawavoluta Noda, 1980:33. (syn. nov.) Type species: (by original designation) Benthovoluta okinav- ensis MacNeil, 1961, Pliocene-Recent. Cyomesus Quinn, 1981:72,73. Type species: (by original designation) Fasciolaria (Mesorhytis) meekiana Dall, 1889, Recent. Diagnosis: Ptychatractinae with fusiform shell, predom- inantly axial sculpture, early teleoconch whorls shoul- dered at intersection between axial ribs and spiral cord, shoulder cord becoming obsolete on subadult and adult whorls; persistence of axial sculpture on adult whorls var- iable in different species. No labral spine or labral spine fasciole. Aperture high, siphonal canal short, broad. Col- umella with three plaits, the adapical one stronger. Latiromitra cryptodon (P. Fischer, 1882) (Figures 1, 3, 5, 10) Mitra cryptodon P. Fischer, 1882:273. Synonyms: Latiromitra specialis Locard, 1897:321, pl. 14, figs 30-34. Teramachia chaunax Bayer, 1971:198, figs 54 (right), 55B— C. (syn. nov.) Other references: Latiromitra cryptodon: Bouchet & Warén, 1985:255, figs 675, 676. Teramachia chaunax: Rehder, 1972:8; Abbott, 1974:243; Bouchet & Poppe, 1995:504. Benthovoluta chaunax: Cernohorsky, 1973:127. Cyomesus chaunax: Quinn, 1981:74, fig. 3 (holotype illus- trated). Type material: Mitra cryptodon and Latiromitra speci- alis: objective synonyms, based on the same two syntypes in MNHN. Teramachia chaunax: holotype USNM 701216. Type locality: Mitra cryptodon and Latiromitra speci- alis: off Morocco, 33°09’N, 09°38'W, 1900 m [R/V Tra- vailleur 1882, dr. 40]. Teramachia chaunax: off St Lucia, 13°45.5'N, 61°05.7'W, 201-589 m [R/V John Elliott Pillsbury, sta. P-904]. Material examined: The type material. MOROCCO. R/V Hirondelle, sta. 116, 31°43’N, Figure 4 Radulae of Latiromitra spp. Latiromitra crosnieri Bouchet & Kantor, sp. nov. (A—C) (paratype, Madagascar, R/V Vauban, chalutage 22): A. Dorsal view of the radular ribbon. B. Enlarged rachidian teeth. C. Radular teeth at the bending plane of radular ribbon. Latiromitra okinavensis (D—-F) (paratype of B. sakashitai, NSMT-Mo 52581). D. Dorsal view of the radular ribbon. E. Enlarged rachidian teeth and right row of raised lateral teeth. E Damaged rachidian tooth, showing the shape of the basal part. Scale bars 50 um (A, D, E); 20 pm (B, C, F). Figure 5 Latiromitra cryptodon. A, B. Syntypes of Mitra cryptodon (MNHN, shell height 30.0 mm); C. Holotype of Tera- machia chaunax (USNM 701216, shell height 27.9 mm). D. Central Atlantic, Tyro Bank (SEAMOUNT 2, sta. DW276, shell height 26.5 mm). E. Brazil (MD55, sta. CB79, shell height 20.2 mm). E G. Brazil (MD55, sta. CB79, shell height 22.5 mm) (G, shell turned to show the columellar plaits). H. Bahamas (R/V Columbus Iselin, sta. 54, shell height 37.6 mm, UMML 30.8272). I. Bahamas (R/V Columbus Iselin, sta. 309, shell height 43.5 mm, UMML 30.9909). J. Bahamas (R/V Columbus Iselin, sta. 156, shell height 37.5 mm, UMML 30.8265). K. Protoconch (same specimen as E, scale bar 500 wm). The Veliger, Vol. 43, No. 1 Page 9 P. Bouchet & Y. I. Kantor, 2000 Page 10 10°47'W, 2165 m, 1 dd (MOM).—R/V Discovery, sta. 8968, 31°35'N, 11°02’W, 1767-1846 m, 1 lv (BMNH). CENTRAL ATLANTIC. The Azores, R/V Princesse- Alice, sta. 683, 38°20’N, 28°05’W, 1550 m, 1 dd (MOM).—SEAMOUNT 2, Tyro Bank, R/V Suroit, sta. DW276, 34°02'N, 28°19'W, 1520 m, 1 lv (MNHN),. BAHAMAS. R/V Columbus Iselin, sta. 54, 23°54'N, 77°13'W, 1298 m, 1 dd (UMML 30.8272).—Sta. 79, 23°51'N, 76°51’W, 1289 m, 1 dd (UMML 30.8262).— Sta. 156, 23°44.4'N, 76°48.3’W, 1334 m, 1 dd (UMML 30.8265).—Sta. 309, 23°44’N, 76°47’W, 1313 m, 1 dd (UMML 30.9909). BRAZIL. MD55: R/V Marion-Dufresne, sta. CB78, 18°59’S, 37°48'W, 1200 m, 2 dd (MNHN).—Sta. CB79, 19°02'S, 37°48'W, 1500-1575 m, 2 lv [1 dissected], 6 dd (MNHN). Distribution: Off Morocco and the Azores (Bouchet & Warén, 1985), now extended to the Caribbean and SE Brazil (Figure 10), collected alive in 1500-1850 m. Description: Shell slender, fusiform, solid, consisting of 2.5 protoconch and 7.5 teleoconch whorls. Protoconch I with large smooth nucleus, diameter 640 ym (Figure 5K); protoconch II consisting of 2.0 convex whorls, smooth, with a subsutural row of coarse granules and a basal keel that is covered by successive whorls, and one to four opisthocyrt incremental riblets before the protoconch/te- leoconch discontinuity. Diameter of protoconch II about 1320 wm. Teleoconch whorls convex, with slightly ad- pressed suture. Sculpture predominantly axial, consisting of solid orthocline ribs, weaker and slightly opisthocline adapically above shoulder cord. The number of axial ribs increases from ca. 10 on the first to third teleoconch whorls to 18—24 on last adult whorl. First teleoconch whorl with five or six weaker spiral cords; the one at shoulder is stronger and persists until penultimate or last adult whorl; other spiral cords usually become obsolete after second whorl. Last adult whorl with spiral grooves, indistinct above periphery, ca. 20, stronger toward base and siphonal canal, where they delimit spiral cords of equivalent breadth. Aperture ovoid, elongate, ca. 50% of total shell height. Outer lip thin, straight. Siphonal canal broad, long but indistinctly set off. Columella with three The Veliger, Vol. 43, No. 1 plaits, seen when shell is rotated clockwise, adapical one stronger. Color creamy to slightly orange-white, periostracum brownish. Protoconch yellowish. Dimensions: [measure- ments from syntype] height 30.0 mm, last teleoconch whorl height 19.0 mm, aperture height 11.0 mm, siphonal canal height 4.3 mm, diameter 10.0 mm. Largest speci- men, height 55 mm. Remarks: Caribbean specimens identified as Teramachia chaunax reach a larger adult size, up to 55 mm, and are more slender than specimens from the NE Atlantic or off Brazil. However, the protoconchs and patterns of teleo- conch sculpture are identical, and there is no doubt that a single tropical Atlantic species is involved. Latiromitra meekiana (Dall, 1889) comb. nov. (Figures 6 B—D, 10) Fasciolaria (Mesorhytis) Meekiana Dall, 1889a: 172, pl. 36, fig. 7; 1889b:112, pl. 36, fig. 7. Other references: Fasciolaria (Mesorhytis) meekiana: Johnson, 1934:127; Rehder, 1972:8. Mesorhytis meekiana: Cernohorsky, 1970:52; 1972:218. Teramachia meekiana: Bayer, 1971:197, figs. 54 (left), 55 D, E; Bouchet & Poppe, 1995:504. Benthovoluta meekiana: Cernohorsky, 1973:127, fig. 2. Cyomesus meekianus: Quinn, 1981:73, 74, fig. 1 (holotype illustrated). Type material: Lectotype, designated by Quinn (1981), USNM 86970. Type locality: Cuba, off Morro Light, La Habana, 732 m [R/V Blake, sta. 100]. Material examined: The lectotype. CARRIBEAN SEA. R/V John Elliott Pillsbury, sta. 1225, 17°42.5'N, 77°58'W, 457-558 m, 1 dd (UMML 30.8260). Distribution: Gulf of Mexico and Caribbean Sea, in 400-730 m (dead collected) (Figure 10). Diagnosis: Protoconch paucispiral, 1.3 whorl, diameter 875 ym. First three teleoconch whorls with strong, broad, Figure 6 Latiromitra aratiuncula (A). A. Holotype (USNM 784594, shell height 20.0 mm). Latiromitra meekiana (B—D). B. Lectotype (USNM 86970, shell height 15.2 mm). C, D. Caribbean Sea (R/V John Elliott Pillsbury, sta. 1225, shell height 26.0 mm, UMML 30.8260—radula and operculum illustrated by Bayer, 1971, figs. 55, D-E). Latiromitra barthelowi (E-J). E, E Holotype (USNM 238444, shell height 27.4 mm). G. Indonesia, Tanimbar Islands (KARUBAR, sta. CC41, shell height 52.0 mm—anatomy see Figure 2A). H. Indonesia, Tanimbar Islands (KARUBAR, sta. CC72, shell height 48.7 mm—radula see Figure 3D-F). I. Indonesia, Tanimbar Islands (KARUBAR, sta. CP59, shell height 38.2 mm), shell turned to show the columellar plaits. J. Protoconch (KARUBAR, sta. CP70, scale bar 500 wm). ?Latiromitra styliola (K—M). K. Lectotype (USNM 108440, shell height 6.5 mm). L. Paralectotype (USNM 880282, shell height 6.6 mm). M. Paralectotype (USNM 880282, shell height 8.5 mm). S) S So N w fe) — S fo} M = > & — iD) re 1S) =) © aa A Page 12 axial ribs, 10 per whorl, subsequent whorls smooth; three ill-defined spiral cords in subsutural zone, cord at shoul- der stronger, intersection with axial ribs angular. Colu- mellar folds 3. Color white. Maximum dimensions 26.0 mm. Remarks: L. meekiana differs from L. cryptodon by its paucispiral protoconch, last teleoconch whorl without ax- ial sculpture, and proportionally higher aperture (63% of total shell height vs. 51% in cryptodon). Latiromitra aratiuncula (Quinn, 1981) comb. nov. (Figures 6A, 10) Cyomesus aratiunculus Quinn, 1981:75, fig. 4. Type material: Holotype USNM 784594. Type locality: Off Virgin Islands, 18°26.4'N, 63°12.6'W, 430 m [R/V John Elliott Pillsbury, sta. P-984]. Material examined: The holotype. Distribution: Off Virgin Islands, 430 m (dead collected) (Figure 10). Diagnosis: A slender, coarsely sculptured Latiromitra species, height/diameter 3.1, last whorl occupying 66% of total shell height. Sculpture consisting of strong axial ribs, axial extension restricted to whorl periphery, 13 on penultimate whorl, fading out in last part of last whorl; raised spiral cords, with narrower interspaces, covering whole exposed height of whorls, nine on antepenultimate and penultimate whorls, 41 on last adult whorl extending onto base and canal. Siphonal canal long. Columellar folds 3. Color probably white. Maximum dimensions (ho- lotype) 20.0 mm, diameter 9.3 mm. Remarks: The holotype and only known specimen is worn and imperfect. The protoconch is unknown, and the early teleoconch whorls are very worn. However, L. ar- atiuncula is a very distinctive species, the only one with ribs restricted to short axial extensions near whorl pe- riphery, and spiral cords covering whole whorl height. ?Latiromitra styliola (Dall, 1927) comb. nov. (Figures 6K—M, 10) Mitra styliola Dall, 1927:48. Other references: Costellaria styliola: Kaicher, 1974: card 266. Type material: Lectotype (designated by Kaicher, 1974) USNM 108440 and 10 paralectotypes USNM 880282. Type locality: Off Georgia, 30°44’N, 79°26'W [R/V AI- batross, sta. 2415]. In the original description, the depth is given as 440 fathoms and 948 meters, which do not match each other (440 fms = 805 m). Material examined: The lectotype and paralectotypes. The Veliger, Vol. 43, No. 1 Distribution: Off Georgia, southeastern United States (Figure 10). Diagnosis: A small, coarsely sculptured Latiromitra spe- cies, height/diameter 2.05—2.21, last whorl occupying 68— 69% of total shell height. Protoconch paucispiral, con- sisting of 1.5 smooth whorls, protoconch/teleoconch dis- continuity sharp. Sculpture consisting of strong axial ribs, reaching shell base, 12 on penultimate whorl, 13 on the last whorl (of 8.5 mm high paralectotype); raised spiral cords, with narrower interspaces, covering whole exposed height of whorls, six on antepenultimate and penultimate whorls, 20 on last adult whorl (of 8.5 mm high paralec- totype) extending onto base and canal. Siphonal canal moderately long. Columellar folds 3. Color white. Max- imum dimensions (paralectotype) 10.3 mm. Remarks: The lectotype and all paralectotypes are dead and worm young specimens that bear a general resem- blance to species such as L. aratiuncula and L. cryptodon. In particular, the stronger spiral cord in the adapical third of the exposed whorl height and the fine, even basal cords on the last whorl appear to suggest a position in Latirom- itra rather than in Mitroidea. However, the lack of pre- served material and even adult shells does not allow us to draw firm conclusions, and we do not exclude the pos- sibility that the species actually belongs to the family Costellariidae as suggested by Kaicher (1974). Mitra sty- liola has been illustrated only once before, by Kaicher (1974: card 266) who figured one of the 11 syntypes as “the type’’, which constitutes a lectotype designation un- der article 74b of the Code of Nomenclature. An appli- cation to place Kaicher’s work on the Official Index of Rejected Works has been rejected by the International Commission on Zoological Nomenclature (Opinion 1905). Latiromitra barthelowi (Bartsch, 1942) (Figures 2A—C, 6E-J, 11) Prodallia barthelowi Bartsch, 1942:12, 13, pl. 2, fig. 2. Other references: Teramachia barthelowi: Weaver & DuPont, 1970:177, pl. 75C, D (holotype illustrated); Bayer, 1971:196—-198; Bouchet & Poppe, 1995:504. Benthovoluta barthelowi: Rehder, 1972:7; Cernohorsky, 1973:127. Cyomesus barthelowi: Quinn, 1981:76, fig. 5 (holotype il- lustrated). Latiromitra barthelowi: Bouchet & Warén, 1985:255, fig. 390. Type material: Holotype USNM 238444. Type locality: Philippines, Sulu Sea, off Cagayan Island, 09°37'45"N, 121°11’E, 905 m [R/V Albatross, sta. 5425]. Material examined: The type material. INDONESIA, TANIMBAR ISLANDS. KARUBAR: R/V Baruna Jaya 1, sta. CP39, 08°36'S, 131°33’E, 466— P. Bouchet & Y. I. Kantor, 2000 Page 13 477 m, 1 lv.—Sta. CC41, 07°45’S, 132°42’E, 401-393 m, 1 lv.—Sta. CC56, 08°16'S, 131°59’E, 552-549 m, 2 lv, 1 dd.—Sta. CP59, 08°20’S, 132°11’E, 405-399 m, 4 lv, 2 dd.—Sta. CP69, 08°42’S, 131°53’E, 356-368 m, 1 dd.— Sta. CP70, 08°41’S, 131°47’E, 413-410 m, 1 lv.—Sta. CP71, 08°38’S, 131°44’E, 477-480 m, 1 juv. lv.—Sta. CP72, 08°36’S, 131°33’E, 699-676 m, 2 lv.—Sta. CP75, 08°46’S, 131°36’E, 452-451 m, 2 lv.—Sta. CP91, 08°44’S, 131°05’E, 884-891 m, 1 dd. INDONESIA, KAT ISLANDS. KARUBAR: R/V Ba- runa Jaya 1, sta. CC21, 05°14'S, 133°00’E, 688-694 m, 1 dd. PHILIPPINES. MUSORSTOM 2: R/V Coriolis, sta. CP79, 13°44’N, 120°32'E, 682-770 m, 1 lv.—MUSOR- STOM 3: R/V Coriolis, sta. CP122, 12°20'N, 121°42’E, 673-675 m, 1 dd. (all MNHN). Distribution: Philippines and eastern Indonesia, alive in 401-682 m (Figure 11). Diagnosis: A large Latiromitra, height/diameter 2.7—3.0 (adults) to 2.95—3.1 (subadults), last whorl occupying 60— 68% (average 62%, n = 7) of total shell height, 52.0 mm high at 8.0 teleoconch whorls. Protoconch (Figure 6J) paucispiral, of 1—1.3 smooth whorls, diameter 1100-1150 pm. First five teleoconch whorls with 14-18 strong rounded axial ribs, interspaces narrower than ribs, sub- sequent whorls generally smooth, but axial ribs occasion- ally persistent until seventh whorl. Spiral sculpture with shoulder cord, defined by its abapical margin, usually dis- tinct on first three to five whorls, but sometimes obsolete; last whorl occasionally with a few low, indistinct narrow cords, but usually smooth; eight to 12 broad cords on base and canal, interspaces much narrower than cords. Siphonal canal rather short. Columellar folds 3, abapical one very weak, sometimes indistinct. Color white, colu- mellar region pinkish, with rather thick, opaque, olive- brown periostracum. Maximum dimensions 52.0 mm, di- ameter 17.7 mm. Remarks: Despite its small size (height 27.4 mm), the holotype from the Philippines appears to be adult, based on its slightly flaring outer lip. It is considerably narrower (h/d = 3.4) than any other specimen, and its high, broad, flat-topped axial ribs are not shouldered by a spiral cord. It contrasts with the Indonesian material, which is rather homogeneous but comes from a restricted geographical area. The MUSORSTOM specimens from the Philippines have rather intermediate characters, and therefore we be- lieve that all the material examined is conspecific. Latiromitra paiciorum Bouchet & Kantor, sp. nov. (Figures 7 A-C, 11) Type material: Holotype and three paratypes in MNHN. Type locality: New Caledonia, off Cape Bayes, 21°05’S, 165°50’E, 960-1100 m [BATHUS 1, sta. CP661]. Material examined: NEW CALEDONIA. BATHUS 1: R/V Alis, sta. CP661, 21°05’S, 165°50’E, 960-1100 m, 1 lv, 3 dd (holotype, three paratypes). BIOCAL: R/V Jean-Charcot, sta. CP61, 24°11’S, 167°32’E, 1070 m, 1 dd. Distribution: New Caledonia and Norfolk Ridge (Figure 11). Description (adult holotype): Shell slender, fusiform, solid, consisting of 2+ protoconch and 7.5 teleoconch whorls. Protoconch I and initial part of protoconch II missing, remaining part of protoconch II broadly conical, with smooth, moderately convex whorls, a few opistho- cyrt axial ribs before the sharp protoconch/teleoconch boundary. Teleoconch whorls convex, slightly concavely depressed in subsutural ramp of last two whorls, suture impressed. Sculpture predominantly axial, consisting of solid orthocline ribs, 10 on first whorl, nine on whorls 2— 3, increasing to 12 on penultimate whorl, ribs generally connected axially to those of preceding and successive whorls, on last adult whorl forming only weakly defined undulations. One much weaker but distinct spiral cord situated at about one-third of exposed whorl height below adapical suture, its adapical margin indistinct, abapical margin abruptly elevated, forming asymmetrical bead at intersection with axial ribs; other spiral cords much more weakly defined, one or two in subsutural ramp, two or three below main cord. On penultimate whorl, the shoul- der cord becomes less sharply defined and there are nine equally weak spiral cords; on last whorl, these spiral cords are still more weakly defined, more broadly spaced near periphery, and there are 25 well-defined, evenly spaced cords on base and canal, interspaces narrower than cords. Aperture narrow, ovoid, elongate, less than 50% of total shell height. Outer lip thin, straight, regularly con- vex, inner lip with a thin opaque callus over parietal area. Siphonal canal broad, short. Columella with three strong plaits, adapical one stronger. Color light brown-beige, polished, periostracum transparent. Protoconch brown. Dimensions of holotype (the largest specimen known): shell height 25.1 mm, last teleoconch whorl height 14.6 mm, aperture height 10.0 mm, siphonal canal height 2.4 mm, shell diameter 8.7 mm. Remarks: The better preserved protoconch of a paratype has 3.2 whorls (Figure 7C), but the initial part (probably only the nucleus of protoconch J) is missing. It has a sculpture of one strong subsutural cord, and there are about eight opisthocyrt axial ribs before the protoconch/ teleoconch boundary, more closely set just before the end of the protoconch. The teleoconch sculpture of the para- types differs from that of the holotype in that the axial ribs extend onto the last whorl. Latiromitra paiciorum resembles L. cryptodon in gen- eral appearance, but differs in having fewer axial ribs (10-14 vs. 18—24 on last adult whorl), stronger columel- Page 14 The Veliger, Vol. 43, No. 1 Figure 7 Latiromitra paiciorum Bouchet & Kantor, sp. nov. (A—-C). A. Holotype (MNHN, shell height 25.1 mm). B. Paratype (MNHN, shell height 24.6 mm). C. Protoconch (same specimen as B, scale bar 500 pm). Latiromitra delicatula (D, E, southwest of Taiwan, MNHN, shell height 33.3 mm). Latiromitra cacozeliana Bouchet & Kantor, sp. nov. (F—H). F Holotype (MNHN, shell height 33.3 mm). G. Paratype (off Epi, MUSORSTOM 8, sta. CP1051, shell height 59.5 mm). H. Protoconch (paratype, off Erromango, MUSORSTOM 8, sta. CP992; same magnification as C). P. Bouchet & Y. I. Kantor, 2000 lar plaits, and a more multispiral protoconch (3.2 vs. 2.5 whorls). It differs from other Indo-Pacific species by its small adult size and multispiral protoconch. Etymology: Named after the Kanak linguistic group oc- cupying the east coast of New Caledonia near Cape Bayes (the type locality), speakers of the Paici language. Latiromitra cacozeliana Bouchet & Kantor, sp. nov. (Figures 7F—H, 11) Type material: Holotype and two paratypes in MNHN. Type locality: Vanuatu [formerly New Hebrides], off Tanna Island, 19°24’S, 169°29’E, 536-556 m [MUSOR- STOM 8, sta. CP975]. Material examined: VANUATU. MUSORSTOM 8: R/V Alis, sta. CP975, 19°24’S, 169°29’E, 536-556 m, 1 dd (holotype).—Off Erromango, Sta. CP992, 18°52’S, 168°55’E, 748-775 m, 1 dd (paratype; protoconch pho- tographed).—Off Epi, Sta. CP1051, 16°37’S, 168°00’E, 555-558 m, 1 dd (paratype). Distribution: Vanuatu archipelago, 536-775 m (dead) (Figure 11). Description (subadult holotype): Shell slender, fusi- form, solid, consisting of 1.3 protoconch and eight teleo- conch whorls. Protoconch light brown, diameter 825 wm, with large nucleus, subsequent whorl very convex, smooth, with four moderately opisthocyrt axial ribs be- fore the sharp protoconch/teleoconch boundary. Teleo- conch whorls rather flat, suture shallow, moderately ad- pressed. Sculpture predominantly axial, consisting of broad, solid, orthocline ribs, interspaces slightly narrower than ribs, 11 on first whorl, 10 on whorls 2-3, increasing to 13 on penultimate whorl, ribs not connected axially to those of preceding and successive whorls, on last part of last whorl becoming less well defined. One spiral cord situated at about one-fourth of exposed whorl height be- low adapical suture, its adapical margin indistinct, abap- ical margin abruptly elevated, forming asymmetrical bead at intersection with axial ribs; another narrow spiral thread below periphery, indistinct on spire whorls, well defined on last two whorls. Last whorl with 15 axial ribs, two or three ill-defined cords in subsutural ramp, and 25 cords on base and canal, weak and low at level of parietal region, raised and distinct at level of columella, inter- spaces much narrower than cords. In addition, there are very fine incremental lines and extremely fine spiral stri- ae, best seen with oblique light. Aperture narrow, ovoid, elongate, less than 50% of total shell height. Outer lip (chipped) thin, straight, regularly convex, inner lip with a thin transparent callus over parietal area. Siphonal canal broad, moderately long, rather distinctly set off from ap- erture. Columella with three strong plaits, adapical one stronger, abapical one only weakly indicated. Color light Page 15 chocolate brown, lighter on apical whorls, protoconch light brown. Dimensions: height 33.3 mm, diameter 10.5 mm, last whorl height 20.0 mm, aperture height 11.0 mm, siphonal canal height 4.0 mm. Remarks: The largest paratype reaches 59.5 X 18 mm (last whorl height 37.3 mm, aperture height 19.2 mm, siphonal canal height 8.0 mm). The tip of the spire is broken, and the sculpture of the first few whorls is rather worn, but they appear to have essentially the characters of the holotype. On the penultimate and antepenultimate whorls the axial ribs fade out and do not reach the ab- apical suture, and on the last whorl there are only vague undulations that are reminiscent of the axial sculpture. Latiromitra cacozeliana is a very distinctive species, characterized by its slender (h/d = 3.2—3.3), solid, choc- olate brown shell, large adult size, and rather long si- phonal canal. The morphology of the protoconch, which indicates non-planktotrophic lecithotrophic larval devel- opment, probably with short dispersal stage, also differs from that of its congeners. Etymology: After the Greek cacozelia, meaning a vicious imitation, an allusion to the superficial similarity to spe- cies of Benthovoluta. Latiromitra crosnieri Bouchet & Kantor, sp. nov. (Figures 4A—C, 8A-—E, 12) Type material: Holotype and three paratypes in MNHN. Type locality: Off NW Madagascar, 12°49'S, 48°27’E, 925-975 m [R/V Vauban, chalutage 118]. Material examined: MADAGASCAR. R/V _ Vauban, chalutage 22, 12°27'S, 48°10’E, 700-680 m, three lv (paratypes).—Chalutage 118, 12°49'S, 48°27’E, 925-975 m, Idd (holotype). SW PACIFIC. R/V Alis, MUSOR- STOM 7, sta. DW540, 12°27'S, 177°28'W, 600 m, 1 lv.— Sta. CP552, 12°16’S, 177°28’W, 786—800 m, 1 lv. Distribution: Madagascar and NE of Fiji, 600-800 m (live) (Figure 12). Description (holotype): Shell slender, fusiform, solid, consisting of 1.2 protoconch and eight teleoconch whorls. Protoconch paucispiral, with large initial nucleus, indi- cating non-planktotrophic larval development, diameter 950 wm, protoconch/teleoconch boundary sharp. First three teleoconch whorls convex, subsequent whorls rather flat-sided, suture impressed, slightly channelled. First five whorls with strong axial sculpture, consisting of solid or- thocline ribs, 10, 13, 15 on second, third, and fourth whorl, respectively, interspaces as broad as ribs. On the fifth whorl, the ribs fade out, and subsequent whorls are smooth with closely set incremental lines. Axial ribs in- tersected by a weakly marked, broad spiral cord, that is defined by its sharp abapical margin and situated at about Page 16 The Veliger, Vol. 43, No. 1 Figure 8 Latiromitra crosnieri Bouchet & Kantor, sp. nov. (A-E). A, B. Holotype (MNHN, shell height 41.2 mm). C. Paratype, Madagascar (R/V Vauban, chalutage 22, shell height 30.4 mm—radula see Figure 4 A—C). D, E. NE of Fiji (MUSORSTOM 7, sta. DW540, shell height 49.0 mm). Latiromitra okinavensis (F—L). EK G. Holotype, USNM P. Bouchet & Y. I. Kantor, 2000 Page 17 one-third of exposed whorl height below adapical suture. Spiral cord obsolete on last smooth whorls. Last adult whorl smooth above periphery, below periphery sculp- tured by ca. 25 low, broad cords, stronger toward base and canal, interspaces narrower than cords. Aperture rath- er broad, ovoid, elongate, about 50% of total shell height. Outer lip thin, straight, regularly convex, inner lip with a thin opaque callus over parietal area. Siphonal canal broad, short. Columella with three strong plaits, adapical one stronger. Color of protoconch and early whorls white, gradually darkening to light creamy beige toward adult whorls, periostracum transparent. Dimensions: height 41.2 mm, aperture height 21.0 mm, diameter 14.7 mm. Remarks: Latiromitra crosnieri shares with L. barthe- lowi, L. delicatula, and L. okinavensis the morphology of the protoconch and early teleoconch whorls. It differs by the combination of broad outline (height/diameter 2.6—2.8 vs. 3.18—3.6), high aperture, and numerous, closely set axial ribs on early teleoconch whorls. Etymology: Named after Alain Crosnier, then of Centre ORSTOM de Nossi-Bé, who collected the type material in 1972-74 during surveys of shrimp stocks off the coast of Madagascar. Latiromitra delicatula (Shikama, 1971) comb. nov. (Figures 7D, E, 12) Benthovoluta delicatula Shikama, 1971:32, pl. 3, figs. 17— 20. Other references: Benthovoluta delicatula: Habe, 1976:97, pl. I, fig. 2; Mat- sukuma, Okutani & Habe, 1991:178, pl. 59, figs. 12, 13}, Type material: Holotype said to be in the Geological Institute, Yokoyama National University (not seen). Type locality: South China Sea, depth unknown. Material examined: Southwest of Taiwan, 1 dd (MNHN ex coll. Poppe). Distribution: Known from South China Sea and off Tai- wan (Figure 12). Diagnosis: A slender, narrow Latiromitra species, height/ diameter 3.3—3.6, last whorl occupying 68% of total shell height, 33.2 mm high at 6.5 teleoconch whorls. Proto- conch paucispiral, of 1.3 smooth whorls, diameter 950 pm. Eleven strong axial ribs on first four teleoconch whorls, fading out on subsequent whorls, last adult whorl smooth. Spiral sculpture with one well-defined shoulder cord and four to six additional, very weakly defined cords on first four spire whorls, only abapical margin of shoul- der cord perceptible on subsequent whorls; about 12 cords on base and canal. Siphonal canal long. Columellar folds 2, deep inside aperture. Color white. Maximum di- mensions (holotype) 43.8 mm, diameter 12.3 mm. Latiromitra okinavensis (MacNeil, 1961) comb. nov. (Figures 4D—F; 8F—H, 12) Benthovoluta okinavensis MacNeil, 1961:96, pl. 9, figs. 2, 3f Synonyms: Benthovoluta sakashitai Habe, 1976:97-—99, pl. I, fig. 3. (syn. nov.) Other references: Okinawavoluta okinavensis: Noda, 1980:33, 34, pl. 7, figs. 16a, b, pl. 10, figs. 20a, b, pl. 11, fig. 8; Noda, 1988: 48, pl. 11, figs. 2a—3b. Type material: B. okinavensis: holotype USNM [Dept of Paleobiology] 562841. B. sakashitai: holotype NSMT- Mo 52579, two paratypes NSMT-Mo 52580, 52581, all live taken. Type locality: B. okinavensis: Okinawa, Chinzato For- mation (Pliocene). B. sakashitai: southern Japan, Kyushu, off Makurazaki, Kagoshima Prefecture, 350—360 m. Material examined: The type material. OKINAWA, Chinzato formation, 1 fragment (USNM 353596). Distribution: Okinawa, Pliocene; Kyushu, Recent, 350— 360 m (alive) (Figure 12). Diagnosis: A slender, narrow Latiromitra species, height/ diameter 3.18—3.86, last whorl occupying 64—68% of to- tal shell height, 47.5 mm high at 7.5 teleoconch whorls. Protoconch paucispiral, of 1.2—1.3 smooth whorls, di- ameter 1000 pm. 10-11 strong axial ribs on first teleo- conch whorls, increasing to 13 on penultimate whorl, fad- ing out on last whorl. Spiral sculpture with one broad shoulder cord, especially strong and well defined on first four whorls, and additional, weakly defined cords; all cords thin, rather indistinct and equal on last two whorls, ca. eight to 10 on penultimate whorl; about 12 cords on base and canal. Siphonal canal long. Columellar folds 2, deep inside aperture. Color brownish flesh. Maximum di- mensions (holotype of B. sakashitai) 47.5 mm, diameter 14.0 mm. 562841, shell height 47.2 mm. H. Fragment of the shell, Okinawa, Chinzato Formation (Pliocene), height of the remaining part 14.6 mm. I-K. holotype of Benthovoluta sakashitai Habe, 1976 (NSMT-Mo 52579, shell height 47.5 mm). L. paratype of B. sakashitai (NSMT-Mo 52581, shell height 47.2 mm—radula see Figure 4D-F). Page 18 The Veliger, Vol. 43, No. 1 Remarks: Comparison of the holotype and published il- lustrations of Latiromitra okinavensis (Noda, 1980, 1988) with the types, and only known material, of Benthovoluta sakashitai demonstrates their conspecificity. Latiromitra okinavensis is distinguished from its congeners by its slender shell with higher last whorl. The most similar Pacific species is Latiromitra barthelowi, which differs in having a less slender shell and more numerous axial ribs. The genus Okinawavoluta was established for this sin- gle species. Noda (1980) compared it with Benthovoluta, Latiromitra being then virtually unused. The shell mor- phology leaves no doubt of it being congeneric with the Recent species placed by us in Latiromitra. Latiromitra vitilevensis (Ladd, 1982) comb. nov. (Figures 9, 12) Volutomitra? vitilevensis Ladd, 1982:56, pl. 14, figs 8, 9. Type material: Holotype USNM [Dept of Paleobiology] 214332. Type locality: Viti Levu, Fiji Is, US Geological Survey station C2026, “‘From bulldozed surface thought to be close to dip slope between C2021 [West of pumping sta- tion Nausori] and C2024 [roadcut west of C2021],”’ 18°02.0’S, 178°31.3’E, Pliocene. Material examined: The holotype. Distribution: Only known from Viti Levu, Fiji; Pliocene (Figure 12). Diagnosis: Shell slender, fusiform, height/diameter 2.90, last whorl occupying 67% of total shell height, consisting of 7.8 teleoconch whorls. Protoconch with only last whorl remaining (probably 0.5+ whorl broken), smooth, with a few opisthocyrt axial riblets before protoconch/teleo- conch discontinuity, diameter 775 wm. Ten strong axial ribs on first teleoconch whorls, increasing to 14 on pen- ultimate whorl, fading out on last whorl. Spiral sculpture with one broad shoulder cord, remaining strong and well defined onto last whorl, and additional, weaker and un- even, but well-defined cords, becoming obsolete abapi- cally on penultimate whorl and on periphery of last whorl; ca. eight on penultimate whorl, four adapically, four abapically of shoulder cord; about 25 cords on base and canal. Siphonal canal long. Columellar folds 3, well exposed due to chipped aperture. Dimensions: height 34.5 mm, diameter 11.9 mm. Remarks: The rather flat teleoconch whorls with strong axial ribs and distinct spiral cord on the shoulder are char- acters shared with species of Ceratoxancus and Latirom- itra, but not with those of Volutomitra. V. vitilevensis does not have the labral tooth or groove that is found in several species of Ceratoxancus, including the type spe- cies, and in this respect is similar to the type and other species of Latiromitra. Based on these shell characters, Figure 9 Latiromitra vitilevensis. Holotype USNM 214332, shell height 34.5 mm. we here transfer Volutomitra? vitilevensis to the genus Latiromitra. In the original description, Ladd (1982) not- ed the ‘“‘somewhat obscure’’ family and generic relations. L. vitilevensis shows overall resemblance to L. cryptodon, from which it differs by having fewer axial ribs. What remains of the broken protoconch suggests that the spe- cies had non-planktotrophic larval development; however, the presence of opisthocyrt axial riblets before the pro- toconch/teleoconch boundary may be correlated with a short lecithotrophic dispersal stage. The associated fauna at the type locality indicates a moderately deep-water assemblage. For instance, Pseu- P. Bouchet & Y. I. Kantor, 2000 Table 1 Conchological and radular characters discriminating the species of Latiromitra. cacozeli- vitilevensis styliola meekiana paiciorum — okinawensis cryptodon delicatula crosniert ana IS) barthelowi aratiuncula 34.5 ~1.5 >10.3 47.5 26.0 55.0 43.8 41.2 20.0 Maximum adult size (mm) m S (oe) mo va) nN 1.3 ioe) N.a. Number of protoconch whorls Maximum diameter of 1125 1000 900 775 875 950 spiral cords on the base 950 smooth broad spiral 825 1150 axial ribs + broad spiral protoconch (wm) Sculpture on the last adult obsolete axial ribs + spiral cords smooth axial ribs + axial ribs + axial ribs + axial ribs + spiral cords spiral cords spiral cords spiral cords cords cords on the base spiral cords whorl n.a. 0.40-0.75 n.a. 0.3 0.3 n.a. Operculum length/aperture height Radula width/aperture Va) nm 1.39 0.9 1.4 height, % Radula length/aperture 17.5 n.a. 17 N.a. height, % Number of rows of radular 110 80 0S 130 n.a. teeth Page 19 domalaxis roddai Ladd, 1982, described from the same fossil locality, has subsequently been synonymized by Bieler (1993:322) with Spirolaxis rotulacatharinea (Mel- vill & Standen, 1903), which is recorded alive at depths of 146-302 m. DISCUSSION Anatomy In comparative remarks on the anatomy of Ceratox- ancus, Kantor & Bouchet (1997) enumerated the char- acters that are shared by Latiromitra, Ceratoxancus, and Benthovoluta: (1) short or very short proboscis, (2) paired proboscis retractors, (3) position of the buccal mass and opening of the radular diverticulum into the buccal cavity at the proboscis base in its contracted position, (4) ventral odontophore retractor passing through the nerve ring, (5) presence of a single accessory salivary gland, (6) large gland of Leiblein, (7) mid-esophagus with well-developed dorsal glandular folds, and (8) small stomach. The general anatomy of studied species of Latiromitra is very similar to that of Ceratoxancus. The only signif- icant difference is that in Latiromitra the salivary glands are fused, while in Ceratoxancus the glands are clearly separate. Mode of Development Planktotrophic larval development can be inferred from the multispiral protoconch of L. cryptodon and L. paiciorum. This is correlated with the broad, disjunct dis- tribution of L. cryptodon, which lives on both sides of the Atlantic and on the mid-Atlantic Ridge, separated from each other by abyssal depths. The protoconch of L. paiciorum suggests that it has a wider range than the two stations off New Caledonia currently indicate. Non-plank- totrophic larval development can be inferred from the paucispiral protoconch of L. meekiana, ?L. styliola, L. barthelowi, L. cacozeliana, L. crosnieri, L. delicatula, and L. okinavensis. The protoconch is partly broken in the only known specimen of L. vitilevensis, and what re- mains suggests that the species had non-planktotrophic larval development. Non-planktotrophy is rather unex- pected in L. crosnieri, which is known from two widely separated localities, off Madagascar and NE of Fiji. This suggests that it may be found at many intermediate lo- calities and/or that its lecithotrophic larva has a free- swimming demersal phase. The protoconch is unknown in L. aratiuncula. Composition of the Genus The radular morphology of Latiromitra cryptodon (Ta- ble 1) appears to be very similar to that of Fasciolaria meekiana Dall, 1889, the type species of Cyomesus Quinn, 1981. The radula illustrated by Bayer (1971: fig. 55B, D, as Teramachia chaunax) has a multicuspid rach- The Veliger, Vol. 43, No. 1 @® @ = Latiromitra cryptodon (type locality and examined material) A A Latiromitra meekiana_ (type locality and examined material) © Latiromitra aratiuncula (type locality) = = ?Latiromitra styliola (type locality) Figure 10 Distribution of the Atlantic species of Latiromitra. idian, with three cusps on both sides of the larger central cusp, whereas in another specimen the radula has a nor- mal three-cusped rachidian (Harasewych, 1987). Thus this character seems to be intraspecifically variable or, more likely, the radula illustrated by Bayer may have had a number of teratological radular rows. The radulae of L. barthelowi, L. okinavensis, and L. crosnieri are also sim- ilar to that of L. cryptodon, both in terms of teeth shape and relative dimensions. It should be mentioned that the radula of L. crosnieri is intermediate in terms of rachidian shape between L. cryptodon on the one hand, and L. bar- thelowi and L. okinavensis on the other. At the same time, the radula of L. crosnieri is significantly narrower than that of other studied species. The shell of Latiromitra cryptodon is very similar to the shell of the type species of Cyomesus, as well as other species originally included by Quinn (1981) in Cyomesus: Teramachia chaunax, Cyomesus aratiunculus, and Pro- dallia barthelowi. These shells are characterized by hav- ing a sculpture with strong axial ribs and flattened and evenly convex teleoconch whorls. Therefore there seems to be no reason to consider Cyomesus distinct from La- tiromitra, and we confirm the synonymy established by Bouchet & Warén (1985). In addition to the species already discussed, Benthov- oluta delicatula Shikama, 1971, and B. okinavensis Mac- Neil, 1961, from off Taiwan and Japan respectively, are conchologically similar and should be included in Lati- romitra, as suggested by Harasewych (1987) [as Cyome- sus] (the latter species under the name B. sakashitai). The radula is known only for L. okinavensis, and in all details it is similar to that of other species studied. The same applies for Volutomitra? vitilevensis Ladd, 1982, which is here included in Latiromitra. Latiromitra species differ from Ceratoxancus in having narrower, more lightly built shells, with predominantly axial sculpture, and spiral cords generally limited to the shoulder of the apical whorls and the base of the last P. Bouchet & Y. I. Kantor, 2000 Page 21 © @ Latiromitra barthelowi (type locality and examined material) A A Latiromitra paiciorum (type locality and examined material) @ Latiromitra cacozeliana (type locality and examined material) Figure 11 Distribution of Latiromitra barthelowi, L. paiciorum Bauchet & Kantor, sp. nov., and L. cacozeliana. Bouchet & Kantor, sp. nov. whorl. They also lack a labral spine or fasciole. However, in view of the small anatomical difference between the two genera, recognition of Latiromitra and Ceratoxancus as valid genera may have to be re-evaluated in the future. Ceratoxancus basileus, which lacks a labral fasciole and has obsolete axial sculpture on the last adult whorl, strongly resembles species of Latiromitra. It was placed in Ceratoxancus based on anatomical characters (salivary glands not fused). Latiromitra differs from Metzgeria Norman, 1879, by its rather flat whorls and short, broad siphonal canal. Summing up, the genus Latiromitra appears to be widely distributed at bathyal depths at low latitudes in the Atlantic and Indo-West Pacific. Currently, we include the following species in the genus: Latiromitra aratiuncula (Quinn, 1981). Recent, Carib- bean. L. barthelowi (Bartsch, 1942). Recent, Philippines and Indonesia. L. cacozeliana sp. nov. Recent, Vanuatu. L. crosnieri sp. nov. Recent, Madagascar and NE of Fiji. L. cryptodon (P. Fischer, 1882). Recent, tropical and subtropical Atlantic. L. delicatula (Shikama, 1971). Recent, South China Sea and off Taiwan. L. meekiana (Dall, 1889). Recent, Caribbean. L. okinavensis (MacNeil, 1961). Pliocene, Okinawa— Recent, Kyushu. L. paiciorum sp. nov. Recent, New Caledonia. ?L. styliola (Dall, 1927). Recent, southeastern United States. L. vitilevensis (Ladd, 1982). Pliocene, Fiji. Species excluded from Latiromitra Benthovoluta nakayasui Habe, 1976 Benthovoluta nakayasui: Habe, 1976: 98, 99, pl. 1, fig. 1. Cyomesus nakayasui: Harasewych, 1987: 178. Harasewych (1987) transferred Benthovoluta nakayasui Habe, 1976 to Cyomesus. We have not examined the type material from off Taiwan in 150 m, but according to the photograph published in the original description, the holotype is very similar to specimens of Borsonia symbiotes (Wood-Mason & Alcock, 1891) [= B. sub- Page 22 The Veliger, Vol. 43, No. 1 Latiromitra crosnieri (type locality and examined material) Latiromitra delicatula (type locality and examined material) Latiromitra okinavensis (type locality and examined material) Latiromitra vitilevensis (type locality) Figure 12 Distribution of Latiromitra crosnieri Kantor & Bouchet, sp. nov., L. delicatula, L. okinavensis, and L. vitilevensis. corpulenta (E. A. Smith, 1899) = B. ochracea (Thiele, 1925)] from the Philippines and Indonesia in MNHN (A. V. Sysoev, personal communication). In the original description Habe mentioned “‘weak growth lines sinuating below the suture as in the family Turri- dae,” as well as three plaits on the columella. Borsonia symbiotes also has columellar folds, usually a single one, but three folds are present in one of the specimens examined. Neither radula nor operculum was studied by Habe. Therefore we think that Benthovoluta nakayasui should be transferred to the family Conidae [sensu Tay- lor et al., 1993] and probably synonymized with Bor- sonia symbiotes. Fasciolaria (Mesorhytis) costata Dall, 1890 Fasciolaria (Mesorhytis) costatus Dall, 1890:317, pl. 5, figes5! Benthovoluta costata: Cernohorsky, 1973:129. Cyomesus costatus: Quinn, 1981:74, 75, fig. 2. Fasciolaria (Mesorhytis) costata is a small (shell height lematical. The shell has very convex whorls with widely spaced axial folds, also present on the last adult whorl. These differences between F. costata and other species of Latiromitra seem sufficient to exclude it from that genus and place it tentatively in Metzgeria. Vexillum (Latriomitra?) [sic] problematicum Ponder, 1968 Vexillum (Latriomitra?) [sic] problematicum Ponder, 1968: 45, pl. 4, figs. 55, 56. Latiromitra problematica: Dell, 1995: 19, fig. 22. This species from 86-540 m (dead collected) off SE New Zealand was tentatively attributed by Ponder to the sub- genus Latiromitra, since the radula was unknown. Dell (1995) reported 14 additional dead collected specimens from Papanui Canyon, Otago, and off the Antipodes Islands. The shell has very convex whorls with widely spaced axial ribs, suggesting a position in Metzgeria rather than Latiromitra. 13.6 mm) deep-water Caribbean species known only Acknowledgments. M. G. Harasewych, R. Germon, W. Blow, from its dead holotype collected in 1256 m. The inclu- and T. Waller (USNM), Hiroshi Saito (NSMT), Bruce Marshall sion of the species in Cyomesus (= Latiromitra) is prob- (NMNZ), and Nancy Voss (UMML) arranged for the loan of P. Bouchet & Y. I. Kantor, 2000 material under their care. We also would like to thank Bruce Marshall for continued support of this project, from processing expedition material as a visiting curator in MNHN to reading and commenting on the manuscript. Gary Rosenberg (Academy of Natural Sciences, Philadelphia) drew our attention to Mitra styliola as a putative member of the Turbinellidae. The junior author wants to express his thanks to the staff of MNHN for their kind assistance during work in the museum, especially to Vir- ginie Heros, Philippe Maestrati, and Pierre Lozouet. The work was carried out during a stay of the junior author as visiting curator in the Muséum national d’ Histoire naturelle, Paris. LITERATURE CITED ABBOTT, R. T. 1974. American Seashells. 2nd ed. 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Bulletin of Zoological Nomenclature 55(3):191—193. PONDER, W. FE 1968. Nomenclatural notes on some New Zealand rachiglossan gastropods with descriptions of five new spe- cies. Records of the Dominion Museum 6(4):29—47. QuInn, J. F 1981. A new genus of Turbinellidae (Gastropoda: Prosobranchia), with the description of a new species from the Caribbean Sea. The Nautilus 95(2):72-77. REHDER, H. A. 1972. Some notes on the genus Teramachia (Vol- utidae: Calliotectinae). The Veliger 15(1):7—10. SHIKAMA, T. 1971. On some noteworthy marine Gastropoda from Southwestern Japan (III). Science Reports of the Yokoyama National University. Section II, Biology—Geology, 18:27—35. TayLor, J. D., Yu. I. KANToR, & A. V. Sysogv. 1993. Foregut anatomy, feeding mechanisms, relationships and classifica- tion of Conoidea (= Toxoglossa) (Gastropoda). Bulletin of the Natural History Museum, London, Zoology 59:125—169. THIELE, J. 1929. Handbuch der systematischen Weichtierkunde. Teil. 1. Gustav Fischer: Jena. 376 pp. WEAVER, C. S. & J. E. Dupont. 1970. Living volutes. A mono- graph of the Recent Volutidae of the World. Delaware Mu- seum of Natural History, Monograph Series, 1:1—375. The Veliger 43(1):24—33 (January 3, 2000) THE-VELIGER © CMS, Inc., 2000 Sex Change, Reproduction, and Development of Crepidula adunca and Crepidula lingulata (Gastropoda: Calyptraeidae) RACHEL COLLIN Committee on Evolutionary Biology, University of Chicago, Culver Hall, Rm. 402, 1025 E. 57th Street, Chicago, Illinois 60637, USA (e-mail: rcollin@ midway.uchicago.edu) Abstract. This paper describes the reproduction and development of Crepidula adunca and Crepidula lingulata from San Juan Island, Washington, USA. Both species were sampled every 2 months to determine the reproductive season and to measure the relationship between adult size and sex, capsule number, egg number, and brood weight. Development of each species is described briefly based on the broods obtained from these samples. As expected for protandrous hermaphrodites, females were larger than males, in both species. Crepidula adunca is an intertidal species that is usually found in stacks of two animals. This species reproduces all year, and embryos develop directly into crawl-away juveniles. Crepidula lingulata is a solitary subtidal species that produces planktotrophic veliger larvae in the summer. These veligers have distinctive pigmentation on the velum and settle 4 weeks after hatching at a shell length of 750 ~m. Experiments show that associations with conspecifics do influence sex change in C. lingulata: males kept with females were less likely to change sex than those kept alone or with other males. INTRODUCTION Snails in the family Calyptraeidae (Caenogastropoda), slipper limpets (Crepidula Lamarck, 1799), cup and sau- cer limpets (Crucibulum Schumacher, 1817), and hat shells (Calyptraea Lamarck, 1799), are marine, suspen- sion feeding, protandrous hermaphrodites that brood their young (Hoagland, 1977). Calyptraeids’ sedentary life- styles and diversity of developmental modes make them ideal animals with which to investigate the evolution of life histories (Hoagland, 1977, 1984, 1986; Warner et al., 1996); development (Coe, 1949; Hoagland, 1986); and the evolution of sex change (Coe, 1938a, b, 1942; Hoag- land, 1978; Collin, 1995). Crepidula species exhibit a va- riety of “‘social’’ habits: some species form semi-perma- nent stacks of many individuals, others make small stacks or pairs, and some do not stack. Life history theory pre- dicts that these different strategies may affect size at sex change, so that at the population level, solitary species have one optimal size at sex change, while those that stack show overlap in the sizes of males and females (Charnov, 1982; Collin, 1995). Additionally, Coe (1938b) stated that stacking species change sex in response to conspecifics while solitary species do not. However, these aspects of Crepidula biology have been examined empir- ically for only a few species (e.g., Warner et al., 1996). Crepidula species are also well known for the great diversity in developmental modes found among closely related species. In fact, there is almost as much variation in developmental mode among species of Crepidula as there is among all caenogastropods: Some species have planktotrophic development, some are lecithotrophic, some develop directly from large eggs, and some develop directly from small eggs by consuming nurse eggs. Basic information on life history, reproduction, and develop- ment has been compiled for 22 species of Crepidula (Hoagland, 1986). There are detailed data on New Eng- land species such as Crepidula fornicata (Linnaeus, 1758), and C. plana Say, 1822 (Ament, 1979; Hoagland, 1986; Conklin, 1897; Franzen & Hendler, 1970; Pechenik & Eyster, 1989); and Coe (1942, 1949) made observa- tions on the natural history of some Californian species. However, little is known about most species outside North America (but see Pilkington, 1974; Gallardo, 1977), or species in the Northeast Pacific and Florida. Despite the fact that development mode is unknown for about half the currently recognized species, it is known for more species of Crepidula than for species in any gastropod genus other than Conus (Kohn & Perron, 1994; A. J. Kohn, personal communication). I studied the life history and development of two Cre- pidula species with different development types from San Juan Island, Washington state. The first aim of the study was to describe the relationship between size, intraspe- cific associations, and sex in these species, and to test the following hypotheses: (1) at the population level, solitary species have one optimal size at sex change, while those that stack show overlap in the sizes of males and females (Charnov, 1982; Collin, 1995), and (2) associations with conspecifics affect sex change in stacking species but not in non-stacking species. The second aim of this study was to measure life history and reproductive characteristics, R. Collin, 2000 Page 25 Table 1 Collection dates and number of animals collected. Collection Number Species date of males C. adunca January 18 137 April 5 64 June 17 74 August 27 90 C. lingulata February 17 97 April 14 60 June 11 70 August 13 88 such as the relationship between female size and fecun- dity, frequency of brooding, duration of development, du- ration of planktonic period, and size at setthement, which are all important parameters in models of life history evo- lution (e.g., Caswell, 1981; Havenhand, 1993; Rowe & Ludwig, 1991). The third aim was to briefly describe the development and embryology of these species. MATERIALS anpD METHODS All snails for this study were collected along the coast of San Juan Island (48°34’N, 123°2’W), San Juan County, Washington, USA, between January and August 1996. Specimens of Crepidula adunca Sowerby, 1825, were collected attached to shells of Calliostoma ligatum (Gould, 1849) from the mid-intertidal zone at Deadman’s Bay on the west side of San Juan Island. All other ani- mals were collected, attached to small rocks, with SCUBA from a depth of 20—30 meters in Shady Cove on the east side of San Juan Island. Animals were sampled every 2 months, as tides and weather permitted. Sampling dates and the number of each species collected are pre- sented in Table 1. Within 2 days of collection, the lengths of all animals were measured to within 0.1 mm with calipers, sex was determined on the basis of presence/absence of the penis and female genital papilla, and their broods were collect- ed and observed live. For C. adunca I counted all the embryos in every capsule whenever possible. However, freshly laid eggs sometimes clumped together in such a way that they could not be counted. For C. lingulata I counted all the embryos in a subset of four arbitrarily selected capsules from each brood (and report the aver- age). In all cases the total number of capsules per brood was recorded. Dry weight was measured for arbitrarily selected adults and egg capsules in which the embryos had not begun to secrete a shell. Because the dry weight of individual eggs was too low to be measured accurately, average egg weight was calculated for C. adunca by di- viding the total capsule weight by the number of eggs in that capsule. Because this measure of ‘“‘egg size”’ includes extra-embryonic material, it more accurately reflects fe- Number Number of Number of Total number of females transitionals juveniles of animals 68 0) 53 258 25 1 28 118 45 6 18 146 68 3 6 167 111 3 2 213 70 0) 4 134 76 1 0) 147 76 1 2 167 male investment per offspring rather than absolute egg size. However, calyptraeid egg capsule coverings are ex- tremely thin and probably contribute little to the total weight of the capsule. Egg diameter of uncleaved eggs and shell length at both hatching and settlement were measured with an ocular micrometer on a compound mi- croscope. I performed an experiment to determine if association with conspecifics effects sex change. Males whose shell length fell within the range of size overlap between males and females were randomly assigned to a beaker with one of three treatments: alone, with another smaller male, or with a female. The water was changed every day and the microalgae Rhodomonas, Isochrysis, and Dunaliella were provided ad libitum. After 3 months the animals were measured and sexed. To determine the duration and frequency of brooding, adult animals were maintained in running seawater tables (at 10—12°C) at Friday Harbor Laboratories. Animals at- tached to glass bowls and the sides of the sea table, through which broods could be observed. Calyptraeid development can be observed without re- moving the embryos from the transparent capsules. En- capsulated embryos were observed with a dissecting mi- croscope, and excapsulated embryos were viewed with a compound microscope using both bright and dark field illumination. In preparation for examination with scan- ning electron microscopy (SEM), embryos were relaxed in 7.5% MgCl, fixed directly in 1% OsO, in distilled H,O, dehydrated in a graded series of EtOH, and dried by transferring them from 100% EtOH to Hexamethyld- isalizine (HMDS). The HMDS was allowed to evaporate, the specimens were mounted on a stub, sputter coated with gold and palladium, and viewed with a JEOL JM35 SEM. Larvae of C. lingulata, the only indirect developer in this study, were reared to metamorphosis. After hatching, larvae were transferred to 0.45 wm filtered seawater in glass custard dishes that were kept partially immersed in running seawater tables at ambient sea temperature (11— 12°C). The initial larval density of approximately 1/mL Page 26 [Re] male WN female C. adunca 60 Number C. lingulata Length (mm) Figure 1 Histograms of the relationship between size and sex for (a) Cre- pidula adunca, (b) C. lingulata. was reduced to around 1/5 mL after 2 weeks. Larvae were fed Isochrysis galbana and Rhodomonas sp. ad libitum, and the water was changed every 2—3 days. When the larvae were 25 days old and had begun to grow a cap- shaped shell, I tested a subset of the most advanced in- dividuals for competence to metamorphose. I exposed five to 10 larvae to either adult conditioned seawater (sea- water in which adult animals were kept for 24 hours), a biofilmed dish (dishes that previously had been kept fully submerged in running seawater for at least a week), or adult shells. The Prussian blue method (W. B. Jaeckle, unpublished method) was used to detect embryonic nutrient uptake. Embryos were carefully excapsulated, incubated in a 1 mg/mL solution of ferretin or iron dextran in seawater for 1—2 hours. They were then relaxed, fixed in 10% for- malin, rinsed, and stained with HCl and potassium fer- Table 2 Results of sex change experiment. C. adunca C. lingulata Treatment changed no change changed no change solitary 0) 19 10 2 both males 2 14 12 1 male and female ] 12 3 V The Veliger, Vol. 43, No. 1 rocyanide solutions. In the presence of iron (from the fer- retin or iron dextran) a blue product is formed. Staining was clearly visible with whole mount light microscopy. Voucher specimens are on deposit at the Field Museum of Natural History in Chicago (Crepidula adunca: FMNH #285002; C. lingulata: FANH #285019) RESULTS Relationship between Size, Sex, and Intraspecific Associations The relationship between shell length and sex for Cre- pidula adunca and C. lingulata is shown in Figure 1. Females are generally larger than males, but there is over- lap between the distributions of male and female lengths: 23.5% of the total size range of reproductive individuals for C. adunca, and 37.5% for C. lingulata. Juveniles (generally < 3 mm) were not included because their shells were often broken when they were removed from the substrate and could not be measured accurately. The distribution of male and female sizes did not vary with sampling date in C. lingulata (one-way ANOVA, males: P > 0.5, n = 315; females: P > 0.5, n = 332). However, in C. adunca male size did vary with date (one- way ANOVA: P < 0.005, n = 363), but female size did not (one-way ANOVA: P > 0.25, n = 206). The overall size distribution did not vary with sampling date for any species (one-way ANOVA, C. adunca: P > 0.15, n = 569; C. lingulata: P > 0.5, n = 647). An analysis using a non-parametric ANOVA produced similar results. The frequency of intraspecific associations varied among the species studied here. Crepidula lingulata sel- dom form stacks (2% of all individuals formed stacks of one male and one female). Crepidula adunca, on the oth- er hand, commonly forms stacks of two individuals: 48% of males and 78% of females occurred in stacks. Only 8.8% of C. adunca stacks included three adult individu- als. In all stacks of three there were two males attached separately to one female. In most cases the stacks ap- peared to be permanent because the male shells fitted tightly on the female shell, often leaving a mark. Sex Change Experiments After 3 months, C. adunca in the sex change experi- ments had no visible shell growth, had not laid eggs, and there was high mortality (about 15%), suggesting that cul- ture conditions were suboptimal. The number of C. adun- ca males that changed sex was too small to determine if there is an effect of conspecifics on sex change (Table 2). Crepidula lingulata, on the other hand, did well under these conditions, showing visible shell growth, and sev- eral produced egg masses. Chi-squared analysis of this data show that there is an effect of treatment (Pearson yx? P = 0.001; likelihood ratio x? P = 0.0005), with males that were paired with females less likely to change sex R. Collin, 2000 Page 27 Table 3 Number and percent of females brooding. January— Species February April C. adunca 45/68 (66%) 14/25 (56%) C. lingulata 1/111 (0.9%) 26/70 (37%) than those that were paired with a male or those that were kept alone. Those that changed sex were larger than those that did not (t-test, P < 0.001), but there was no signif- icant difference in size among treatments (Kruskal-Wallis test P = 0.168). In several of the paired male treatments both males changed sex. Reproduction and Brooding The Puget Sound calyptraeid species display a variety of reproductive seasons. Crepidula adunca appears to brood throughout the year, whereas C. lingulata repro- duces in the summer (Tables 3). The number of brooding C. adunca females did not differ between stacked and solitary individuals (Chi-squared test: x? = 0.07644, n = 206, df = 1, P > 0.5). I examined the effect of female body size on several reproductive parameters, using shell length as a proxy for adult body size. Ordinary least squares regression of the Ln (dry body weight) on the Ln (shell length) shows that shell length is a good predictor of body weight for both C. adunca (b = 2.929, n = 102, R? = 0.92, P < 0.005) and C. lingulata (b = 2.50, n = 121, R? = 0.90, P < 0.005) (Figure 2). I used a type I regression because I am most interested in using shell length as a predictor of reproductive parameters (e.g., Collin, 1995). Reduced Major Axis slopes can be calculated from the data given above and in Table 3. There is no difference between the sexes in the relationship between length and dry weight. The relationship between female length, number of cap- sules, eggs per capsule, and total eggs per brood is sum- marized in Table 3 and Figure 2. All measures of repro- ductive output except egg weight in C. adunca clearly increase with female length. However, female size ac- counts for only 10—40% of the variation in any of these parameters (7° in Table 4). In neither species did the num- ber of eggs per capsule vary with developmental stage. There is considerable variation in the size of embryos within any capsule for C. adunca. There sometimes ap- pears to be as much as a twofold difference in diameter of blastulae within a capsule, and there is also consider- able variation in size of later embryos. Some females also have embryos that are larger than those of other females. Unfortunately, the irregular shape of the embryos makes it difficult to quantify the differences especially among embryos at different stages. June August 26/45 (58%) 46/76 (61%) 30/68 (44%) 47/76 (62%) Descriptions of Development C. adunca Development is synchronous within and among cap- sules. The large eggs, 250-350 microns in diameter, cleave equally and almost synchronously. A small polar lobe formed during the first two cell divisions. It may have been present during subsequent divisions, but the size of the eggs made it difficult to see it with any cer- tainty. The first cleavage division takes about 24 hours; in intact capsules incubated at 12° C the second cleavage follows after 18 hours. Third cleavage is unequal and forms four clear micromeres that are easily hidden in the furrows between the macromeres. Later in cleavage, a mound of small, clear micromeres can be seen at the an- imal pole. After 5 days the pile of micromeres has still not yet begun to grow around the macromeres. Gastru- lation occurs by epiboly, as micromere proliferation con- tinues until the large yellow macromeres are completely covered by small clear cells. Because the embryos are large and opaque, it is diffi- cult to observe the details of development with light mi- croscopy. SEMs of the external morphology of several stages are illustrated in Figure 3 and complement the his- tological analysis of Moritz (1939). Dense patches of cilia around the mouth, in the area of the head vesicle, located dorsally between the tentacles, and on the foot anlage develop before any of these features are clearly visible with light microscopy (Figure 3a). In the living embryos, the cilia covering the head and foot beat toward the mouth. The eyes appear as faint pigmented spots before any of the other external features. A cluster of six to eight embryonic kidney cells are visible on either side of the large ciliated mouth (Figure 3a). Each cell has a dense covering of microvilli (Figure 4). Incubation in ferretin followed by staining with the Prussian blue reactions shows that the embryonic kidneys take up protein but not polysaccharides from the external medium from this stage until the kidney cells disappear, shortly before hatching. The shell first appears over a flattened area on the pos- terior dorsal side of the embryo, while the head and foot anlagen are still small and indistinct. It slowly enlarges until it is a miniature version of the virtually bilaterally symmetrical adult shell. At no time is there a coiled em- bryonic shell. One of the most conspicuous characteris- tics of the mid-stage embryo, the head vesicle, a trans- Page 28 Ln Dry Body Weight (g) Ln Total Eggs Crepidula adunca 12S 2 2:5 2.9 275 Ln Length 3 3 3.5 Ln Dry Body Weight (g) Ln Total Eggs 3.25 Figure 2 The Veliger, Vol. 43, No. 1 Crepidula lingulata 15 2 2.5 Ln Length (mm) 20 2.75 Ln Length 3 3 3.5 3.25 Scatter plots of the relationship between shell length and dry weight and shell length and total number of embryos per brood for Crepidula adunca and C. lingulata. Species C. adunca C. lingulata Table 4 Relationship between shell length and reproductive parameters. variable total eggs number of capsules average eggs/capsule average egg weight total eggs capsule number eggs/capsule average weight/capsule mean SD 64.56 29.4 7.14 2.0 8.78 2.48 0.06 mg 0.25 4283 2019 13.56 4.682 281.2 88.93 0.55 mg 0.20 Ordinary Least Squares Regression Statistics of Ln(reproductive parameters) on Ln (length) A n m Ts 85 2.96 0.413 105 1.46 0.284 83 1.42 0.308 17 = 0:32 0.002 81 1.87 0.295 107 0.94 0.101 81 1.35 0.370 63 E22 0.215 P < 0.000001 < 0.000001 < 0.00001 > 0.5 < 0.00001 < 0.001 < 0.00001 < 0.005 R. Collin, 2000 Table 5 Comparative life history data. Length (wm) at Brooding Hatching Planktonic settlement mean Development Egg size Egg number Reproductive Stack- period (SD) duration size mean (SD) type season (wm) 262-315 ing Species 1.5—2.7 mm 275-363 wm ~120 days direct 64.8 (29.3) 4283 year round summer C. adunca 745.2 (82.4) 29-45 days 24-33 days planktotrophic (2020) n C. lingulata Page 29 parent, densely ciliated, fluid-filled sac extends from the head between the tentacles. The head vesicle shrinks (Fig- ure 3c) and disappears before hatching. A small ciliated ridge, the highly reduced velum, is present around the base of each tentacle (Figure 3). The foot differentiates slowly from a ventral bump and at no time is there an operculum. The shell develops pigment before it has reached its full size, and the body also becomes pig- mented at this time. The embryos hatch when all the yolk has been absorbed. The embryo differentiates directly into the juvenile body without displaying any larval or embryonic specializations other than the head vesicle and embryonic kidneys. My observations of the development of C. adunca agree with the few observations of C. adun- ca by Conklin (1897), who focused on the early devel- opment of C. fornicata. The number of embryos per capsule is independent of developmental stage. Embryos hatch without assistance from their mother at shell lengths of 1.5—2.7 mm. Hatch- ling shell length differed significantly between two broods for which I had sample sizes sufficient to estimate statistical parameters (mean = 2.6 mm, n = 11, SD = 0.17 and mean = 1.8, n = 13, SD = 0.10; t-test P < 0.05), and the lengths for a third brood were 1.76—1.98 mm. A mean hatching length of 2.6 mm is unusually large for this species. Hatchlings have no osphradial fil- aments, but the gills, food pouch, and radula are well developed, and pigmentation is the same as the adults. Hatchlings are able to collect and ingest suspended algal cells from the water column, and juveniles did not use their radulae to feed from the substrate. I could not determine the duration of incubation from single broods because females attached to glass did not lay eggs. However, broods kept in dishes at 12° C took about 4 months to hatch. C. lingulata The embryonic development of C. lingulata is the same as the development of C. fornicata described by Conklin (1897) and, in fact, SEM micrographs of C. lin- gulata and C. fornicata embryos are indistinguishable. The eggs are 150 pm in diameter, there are no nurse eggs, and all the embryos within a sac developed synchro- nously. Cleavage proceeds as in C. adunca. Gastrulation is by epiboly with some invagination (Figure 5); the gas- trula is flattened, unlike those of the direct developing species, in which it remains spherical. The embryo grad- ually elongates along the animal-vegetal axis (Figure 6) and the foot, head vesicle, velum, and mouth form as in the other species. It is difficult to distinguish these fea- tures with light microscopy but they are clearly visible in SEMs (Figure 6). Unlike those with direct development, the embryonic kidneys of C. lingulata are made up of only a single cell located on each side of the embryo (Figure 6), and the operculum develops in synchrony with the shell. Page 30 The Veliger, Vol. 43, No. 1 Figure 3 Development of Crepidula adunca: A. SEM of an early embryo in which little structure would be visible with light microscopy. The extensive ciliation over the anterior of the embryo beats into the large mouth. The tentacles are visible on either side of the mouth, and the head vesicle is beginning to bulge between them. The embryonic kidneys are visible to the left of the mouth, and the foot anlage is beginning to differentiate and has a distinctive medial stripe of cilia. The posterior constriction (bottom right) marks the area that is covered by the shell at this stage B., C. Light micrograph and SEM of a later stage in which the head vesicle is well developed, as are the shell and foot. The large visceral yolk mass does not extend into the head vesicle in B, and the head vesicle has collapsed in C. The reduced velum is visible in C. around the base of the tentacles; the ciliation is not distinct from the overall ciliation of the head vesicle and mouth. The embryonic kidneys are visible behind the velum. f, foot; hv, head vesicle; k, embryonic kidneys; m, mouth; sh, shell; t, tentacle; v, velum. Scale bar = 100 microns. Broods laid in the lab took between 24 and 33 days to hatch at 11—-12°C. Those that took longest were laid in spring, while those with the shortest development time were laid later in the summer. Individual females pro- duced at least three broods during the summer, and it took Figure 5 Figure 4 Scanning electron micrograph of a Crepidula lingulata gastrula, Scanning electron micrograph of Crepidula adunca embryonic showing the vegetal cross furrow and the slight invagination as the kidney showing the dense microvilli. Scale bar = 10 microns. micromeres grow around the macromeres. Scale bar = 5O microns. R. Collin, 2000 Figure 6 Light micrographs (A, B, C) and SEMs (D, E, F) of three developmental stages of Crepidula lingulata. A and D show early embryonic stages in which the beginnings of most major structures are just visible in the SEM, but little can be seen with light microscopy. B and E show a mid-stage embryo with obvious, well-developed head vesicle, embryonic kidneys, foot with operculum (E), and a shell that covers about half the yolk. C is a recently hatched veliger, and F shows an embryo just prior to hatching. The head vesicle and embryonic kidneys have disappeared, and the velar cilia are well developed. f, foot; hv, head vesicle; k, embryonic kidney; m, mouth; op, operculum; sh, shell; v, velum. Scale bar = 100 microns. as few as 5—7 days after one brood hatched for another brood to appear. At hatching, the planktotrophic larvae are about 300 zm in shell length. The embryonic shell is smooth, trans- parent, and planspiral or slightly left-handed. Anatomi- cally they resemble larvae of C. fornicata (Werner, 1955): the eyes, tentacles, velum, operculum, heart, and foot are well developed, and the head vesicle and embryonic kid- neys have disappeared. There are distinctive deep red stripes which run parallel and proximal to the food groove on the anterior and posterior curves of each velar lobe. On the lateral edge of the velum, proximal to the food groove, are up to three opaque white spots. The number of spots varies among larvae within a single brood. The only other pigmentation is the black pigment on the in- testine near the shell apex. As the shell grows, it becomes clearly right-handed and develops widely spaced rows of very fine granular sculpture (Figure 7). After 25 days at 11—-12°C the larvae begin to grow limpet-shaped shells, and the velum shrinks. None of these veligers settled when they were exposed to biofilmed substrates, adult- conditioned seawater, or conspecific shells. However, af- ter 27 days, some larvae did settle when exposed to bio- filmed substrate. Larvae settled at an average shell length of 745.2 pm (n = 10, SD = 82.4). There was consider- able variation in the size of the veligers in any one cul- ture, and animals continued to settle in response to biof- ilmed substrates over the next 18 days, while none settled Figure 7 Scanning electron micrograph of the shell of a 3-week-old Cre- pidula lingulata larva. The arrow marks the embryonic/larval shell boundary. Scale bar = 100 microns. in clean glass dishes. I was not able to determine how much longer the veligers would remain competent to set- tle or if they would eventually settle spontaneously in clean dishes. Fretter (1972) described the details of meta- morphosis for this species. DISCUSSION The effects of the proximity of conspecifics on the size at sex change in Crepidula have long been of interest to biologists (Coe, 1938b; Collin, 1995; Warner et al., 1996). Coe (1938b) and Warner et al. (1996) have shown that sex change in gregarious species like C. fornicata and C. norrisiarum Williamson, 1905, is the result of con- specific associations. Juveniles kept alone often bypass the male phase and differentiate directly into females (Coe, 1938b). Males kept in association with females change sex later than those kept with other males. The sex of solitary species was not thought to be effected by conspecifics (Coe, 1938b). If this is true, then variation in stack composition is expected to result in more overlap in the size distribution of the two sexes in populations of species that form stacks. And solitary species are ex- pected to show a more constant size at sex change, re- sulting in little overlap in size between the sexes (Collin, 1995). However, the results of this study suggest that this is not the case. There is considerable size overlap between males and females of both stacking and non-stacking spe- cies. If anything, the overlap of male and female size in the stacking species is less than in the non-stacking spe- The Veliger, Vol. 43, No. 1 cies. This may be due to the demonstrable effect that conspecifics have on the sex change of C. lingulata. It is possible that individuals that live close to one another have the same effect on their associates’ sex as stack- mates do in stacking species. It is clear from the experi- ments described herein that associations among non- stacking species can have an effect on their sex. Unlike the situation in other invertebrate groups and even in other mollusks, there is large intraspecific varia- tion in egg size in C. adunca. Crepidula adunca eggs observed in this study ranged between 262-315 wm for one female. This size range is close to the egg diameter of 240 ym reported by Strathmann (1987) for Friday Har- bor. However, Hoagland (1986; citing Coe, 1949) and Conklin, (1897) gave an egg size of 400 pm for C. adun- ca in California. Unfortunately, it is difficult to find un- cleaved eggs, and therefore this variation in egg size can- not be partitioned into variation among capsules, females, and geographic regions. Size at hatching is also highly variable in C. adunca. The mean hatchling shell length can differ by as much as 60% between broods. Such var- iation in hatching size is common in gastropods that com- pete for nurse eggs in the egg capsules (Rivest, 1983; Hadfield, 1989), but is uncommon in invertebrates that, like C. adunca, develop to hatching without obvious ex- ogenous food supplies. It is becoming a paradigm in studies of the evolution of development that changes in larval or embryonic func- tion (e.g., feeding to non-feeding, or pelagic to benthic) are correlated with often drastic changes in the mecha- nisms and sequence of developmental events (Emlet, 1991). For example, at the turn of the last century, Lillie (1895) explained the modification of cell sizes in unionid bivalve cleavage patterns as an embryonic “‘adaptation”’ to producing the specialized glochidium larva. Recent studies, focusing mostly on echinoids, have highlighted changes in larval skeleton, cleavage patterns, gastrulation, and ciliation in congeners associated with a shift from feeding to non-feeding development (e.g., Henry et al., 1992; Wray, 1996). Contrary to the pattern observed among echinoderms, early development in Crepidula seems to be conserved in the face of changes in egg size and mode of development. There are few morphological differences between direct and indirect developers that are not the direct result of differences in the amount of yolk or the reduction of larval feeding and swimming structures. However, C. adunca develops multiple embry- onic kidney cells, whereas indirect developers like C. lin- gulata and C. fornicata (Conklin, 1897) have only one cell on each side of the body. Additionally, the operculum does not develop in direct developers (it is absent in all adult calyptraeids). Further studies aimed at determining if other selective pressures such as nurse egg feeding and albumin absorption have produced changes in calyptraeid development will further our understanding of adapta- tions in gastropod development. R. Collin, 2000 Page 33 Acknowledgments. I am grateful to R. Strathmann and the fac- ulty and staff of Friday Harbor Marine Laboratories for their support. B. Pernet cheerfully accompanied me on many cold and wet collecting trips; his humor and assistance were much appre- ciated. Will Jaeckle provided technical advice and encourage- ment. This research was supported in part by an NSF pre-doc- toral fellowship and grants from the Pacific Northwest Shell Club, Sigma Xi, the American Museum of Natural History (Ler- ner Gray), and the Western Society of Malacologists. R. Bieler, B. Chernoff, K. E. Hoagland, J. Slapcinsky, and J. Wise made helpful suggestions on earlier versions of this manuscript. LITERATURE CITED AMENT, A. S. 1979. Geographic variation in relation to life his- tory in three species of the marine gastropod genus Crepi- dula: growth rates of newly hatched larvae and juveniles. Pp. 61-76 in E. Stancyk, (ed.), Reproductive Ecology of Marine Invertebrates. University of South Carolina Press: Columbia. CASWELL, H. 1981. The evolution of “mixed” life-histories in marine invertebrates and elsewhere. American Naturalist 117:529-536. CHARNOV, E. 1982. The Theory of Sex Allocation. Princeton University Press: Princeton, New Jersey 353 pp. Cog, W. R. 1938a. Sexual phases in the gastropod Crucibulum spinosa. Journal of Morphology 63:345—361. Cog, W. R. 1938b. Conditions influencing the change of sex in mollusks of the genus Crepidula. Journal of Experimental Zoology 77:401—423. Cor, W. R. 1942. The reproductive organs of the prosobranch mollusk Crepidula onyx and their transformation during the change from male to female phase. Journal of Morphology 70:501—512. Cor, W. R. 1949. Divergent methods of development in mor- phologically similar species of prosobranch gastropods. Journal of Morphology 84:383—400. COLLIN, R. 1995. Sex, size, and position: a test of models pre- dicting the size at sex change in the protandrous gastropod Crepidula fornicata. American Naturalist 146(6):815—831. CONKLIN, E. G. 1897. The embryology of Crepidula. Journal of Morphology 13:3—226. EMLeT, R. B. 1991. Functional constraints on the evolution of larval forms of marine invertebrates: experimental and com- parative evidence. American Zoologist 31:707—725. FRANZEN, D. R. & G. HENDLER. 1970. Substrate diversity and the taxonomy of Crepidula convexa (Say) (Gastropoda: Proso- branchia). Occasional Papers of University of Connecticut (Biology) 1:281—289. FRETTER, V. 1972. Metamorphic changes in the velar muscula- ture, head and shell of some prosobranch veligers. Journal of the Marine Biological Association of the United Kingdom 52:161-177. GALLARDO, C. S. 1977. Two modes of development in the mor- phospecies Crepidula dilatata (Gastropoda:Calyptraeidae) from Southern Chile. Marine Biology 39:241—251. HADFIELD, M. G. 1989. Latitudinal effects on juvenile size and fecundity in Petaloconchus (Gastropoda). Bulletin of Marine Science 45:369-746. HAVENHAND, J. N. 1993. Egg to juvenile period, generation time, and the evolution of larval type in marine invertebrates. Ma- rine Ecology Progress Series 97:247—260. Henry, J. J.. K. M. KtueG & R. A. Rarr. 1992. Evolutionary dissociation between cleavage, cell lineage and embryonic axes in sea urchin embryos. Development 114:931—938. HOAGLAND, K. E. 1977. Systematic review of fossil and recent Crepidula and discussion of evolution of the Calyptraeidae. Malacologia 16(2):353—420. HOAGLAND, K. E. 1978. Protandry and the evolution of environ- mentally mediated sex change: a study of the Mollusca. Ma- lacologia 17:365—381. HOAGLAND, K. E. 1984. Use of molecular genetics to distinguish species of the gastropod genus Crepidula (Prosobranchia: Calyptraeidae). Malacologia 25(2):607—628. HOAGLAND, K. E. 1986. Patterns of encapsulation and brooding in the Calyptraeidae (Prosobranchia: Mesogastropoda). American Malacological Bulletin 4(2):173-183. Koun, A. J. & E E. Perron. 1994. Life History and Biogeog- raphy: Patterns in Conus. Oxford University Press: New York. 106 pp. Lituir, EF 1895. The embryology of Unionidae. Journal of Mor- phology 10:1—100. Moritz, C. E. 1939. Organogenesis in the gastropod Crepidula adunca Sowerby. University of California Publications in Zoology 43:217-248. PECHENIK J. A. & L. S. EysTrr. 1989. Influence of delayed meta- morphosis on the growth and metabolism of young Crepi- dula fornicata (Gastropoda) juveniles. Biological Bulletin 176:14—24. PILKINGTON, M. C. 1974. The eggs and hatching stages of some New Zealand prosobranch molluscs. Journal of the Royal Society of New Zealand. 4(4):411—431. Rivest, B. R. 1983. Development and the influence of nurse egg allotment on hatching size in Searlesia dira (Reeve 1846) (Prosobranchia: Buccinidae). Journal of Experimental Ma- rine Biology and Ecology 69:217—242. RoweE, L. & D. Lupwic. 1991. Size and timing of metamorphosis in complex life cycles: time constraints and variation. Ecol- ogy 72:413—427. STRATHMANN, M. FE 1987. Reproduction and Development of Ma- rine Invertebrates of the Northern Pacific Coast. University of Washington Press: Seattle. 670 pp. WARNER, R. R., D. L. Fitch & J. D. STANDISH. 1996. Social control of sex change in the shelf limpet, Crepidula norri- siarum: size-specific responses to local group composition. Journal of Experimental Marine Biology and Ecology 204: 155-167. WERNER, B. 1955. Uber die Anatomie, die Entwicklung und Biologie des Veligers und der Viliconcha von Crepidula for- nicata L. (Gastropoda Prosobranchia). Helgolander Wissen- schaftliche Meeresuntersuchungen 5:169—217. Wray, G. 1996. Parallel evolution of nonfeeding larvae in echi- noids. Systematic Biology 45(3):308-322. The Veliger 43(1):34—42 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Shell Growth of Mytilus trossulus Gould, 1850, in Port Valdez, Alaska ARNY BLANCHARD! anD HOWARD M. FEDER? Institute of Marine Science, University of Alaska, PRO. Box 757220, Fairbanks, Alaska 99775-7220, USA (‘e-mail: arnyb@ims.uaf.edu, *feder@ims.uaf.edu) Abstract. Shell growth of intertidal Mytilus trossulus in Port Valdez, Alaska, site of a marine oil terminal, is assessed using descriptive methods and the Gompertz growth model. Age-frequency data indicated that mussels of the young to middle age classes (age classes 1 to 5) were dominant in the mussel beds, and individuals older than 7 years were not common. Percentages of new shell growth calculated by month indicated that most new shell growth occurred during the summer with no growth during winter. Growth was greatest for younger mussels with only a little new shell growth for mussels in age class 7 and above. Gompertz growth curves demonstrated differences between sites within Port Valdez related to environmental differences. The addition of a seasonal component to the Gompertz models indicated a strong seasonal amplitude to growth with maximum growth rates estimated to occur during the month of June. Comparisons of curve parameters indicated reduced growth relative to studies from lower latitudes. It is concluded that mussels in Port Valdez are stressed by exposure to natural environmental extremes in the intertidal region resulting in decreased growth. INTRODUCTION Shell growth of Mytilus species has been extensively doc- umented under a variety of natural and laboratory con- ditions, and the general factors controlling this growth are well described (see review by Seed & Suchanek, 1992). It is generally accepted that the timing of growth cycles in mussels is influenced by seasonal temperature ranges and the availability of food resources (Seed & Suchanek, 1992). In addition to temperature and food availability, important factors influencing shell growth include wave exposure and salinity. While exposure to environmental contaminants can adversely affect shell growth in Mytilus (e.g., Stromgren et al., 1986), variability in growth, as a response to natural environmental factors, may mask sub- lethal impacts on mussels (Seed & Suchanek, 1992). Thus, studies assessing the response of mussels to envi- ronmental contaminants have not focused on shell growth alone. Port Valdez, a subarctic fjord in the northeastern corner of Prince William Sound, Alaska, is the site of the marine oil terminal serving as the terminus of the Trans-Alaska pipeline. Recent evidence indicates that the mussel in Port Valdez is Mytilus trossulus Gould, 1850, rather than M. edulis Linnaeus, 1758, as previously believed (e.g., McDonald et al., 1991; see also Blanchard & Feder, 1997). Some aspects of the biology of M. trossulus from Port Valdez were reported by Feder & Keiser (1980) who suggested that the period of peak shell growth follows the onset of spawning in late spring. Monitoring studies relative to the marine terminal include investigations of the ecology of M. trossulus (see review in Shaw & Ha- meedi, 1988). For example, Blanchard & Feder (1997) summarized the findings of reproductive studies of inter- tidal mussels from Port Valdez. This paper describes shell growth of intertidal M. trossulus from Port Valdez, Alas- ka, at sites within and remote from the boundaries of the marine oil terminal. METHODS The Study Area The study reported here assesses intertidal mussel pop- ulations at three sites in Port Valdez, Alaska: Berth 4 (B4, within the confines of the Alyeska Marine Terminal), Sawmill Spit, (SS, a short distance west of B4 and outside of the marine terminal boundaries), and at Mineral Creek (MC, across the port from B4) from January 1981 to Sep- tember 1982 (Figure 1). These sites were among those considered in the reproductive study of Blanchard & Fed- er (1997). The tidal range is approximately 6 m (Colonell, 1980). The annual air temperature in Port Valdez ranges from approximately —10°C to 25°C, and surface water temperatures range from —2 to 16°C (Blanchard & Feder, 1997). In summer, Port Valdez exhibits a positive estua- rine flow with heavy sediment loads and decreased salin- ity (approaching 0%c) while in winter, freshwater input decreases and estuarine circulation ceases (Colonell, 1980). While all sites experience decreased salinity and increased sediment loads in summer, the MC site is most heavily affected by these physical stresses by virtue of its proximity to Mineral Creek, a glacial stream. Sampling and Laboratory Procedures Mussels were collected monthly at the three sites in 1981 (January—November)—1982 (January—September). A. Blanchard & H. M. Feder, 2000 Shoup Glacier Mineral Creek Port Valdez Sawmill Spit —~., e Mineral Creek Page 35 Valdez cee Ne Marine Terminal Sawmill Creek 0 kilometers Figure | Map of sampling locations in Port Valdez, Alaska. Sampling locations are indicated with a large dot. Three destructive plots were taken by randomly tossing a 0.0156 m? (12.5 X 12.5 cm) sampling frame onto the mussel bed in the mid-intertidal zone. Shells were aged in the laboratory by counting growth annuli which were readily apparent due to cessation of growth in mussels during winter in Port Valdez (Feder & Keiser, 1980). Val- idation of the aging technique was performed in a pilot study where the shells of 100 mussels were notched in the spring and examined 1 year later (Feder & Keiser, 1980). Observations the following year indicated that all mussels had formed a single annulus confirming the va- lidity of annuli for use as an aging tool specifically for this Alaskan mussel population. The age of mussels in this study was estimated as the sum of observable growth rings, not including the shell edge. Total shell length, the maximum distance from the umbo to the shell edge, was measured to the nearest 0.01 mm. New shell growth was measured as the maximum distance from the last age an- nulus to the edge of the mussel shell. Although every effort was made to estimate the age of each mussel, spec- imens with shells too badly eroded to age could not be included in the determination of the age to shell-length relationships. The O age class referred to here applies to those individuals of the settling year class that have un- dergone only one partial growing season and survived through only one winter. Movement of a mussel to the next age class occurs when new shell growth can first be observed, usually in the spring of the year. Data Analysis Age-frequency distributions and mean shell size per age class of mussels from each site are summarized on an annual basis. The mean new shell growth and percent new shell growth for each age class by year are also pre- sented. The percentage of new shell growth is plotted against the proportions of gametic tissues and nutritive tissues after Blanchard & Feder (1997). Shell growth was modeled using a Gompertz growth equation. The Gompertz equation is given by: a at TS) Ly = Liex where L, = the length of mussel at age t, L., is the as- ymptotic maximum growth, k = growth coefficient, t = age of mussel, and ty is the curve’s inflection point (Rat- koskwy, 1990); this is a reparameterization of the usual Gompertz growth model applied to mussels (e.g., Seed & Suchanek, 1992). This curve is appropriate for shell growth that follows a sigmoidal curve but assumes that growth is determinate, an assumption that is often appro- Page 36 priate for slow-growing mussels (Seed & Suchanek, 1992). To account for seasonality, a seasonal component was added to the Gompertz growth model resulting in: CK aT ktt ly) — —Sin(2n(t — t.)] L, = Lie 2a where C is the amplitude of the seasonal growth oscil- lation (0 = C < 1), t, estimates the fraction of the year elapsing before the maximum growth rate occurs (see Sparre et al., 1989), and t, (t, + 0.5) indicates the time of the year when growth is minimal. The parameter C gives indication of the strength of the seasonal pattern, as values near zero suggest little change in growth over a year, while values near one indicate maximal amplitude to growth patterns. When C equals one, no growth occurs at time t,. Based on preliminary analyses, the parameter C was fixed at one for all seasonalized Gompertz models. The age of mussels was calculated as the age estimated by growth annuli plus the fraction of the year correspond- ing to the month the sample was taken. For example, an age 4.5 mussel is a mussel with four growth rings col- lected in June (4 + 6/12). To increase the number of specimens in the older age classes, data from 1981 and 1982 were combined by site. Growth model parameters were estimated by the non- linear regression procedure in SYSTAT 7.0 (SPSS Inc., 1997) using the Gauss-Newton maximization algorithm with the least-squares loss function [Loss = (Observed — Expected)’]. Since the sample size for each data set was large (n = 3600), normal confidence intervals were cal- culated for each parameter (95% Cl = parameter + 1.96% Asymptotic Standard Error). Convergence, the ability of the maximization algorithm to calculate the parameter es- timates, is useful to assess the fit of nonlinear models. For models that fit data well, a program will require fewer iterations to determine parameter estimates with only small changes in estimated values in the final iterations. For the Gompertz models applied here, convergence was usually gained in about 10 iterations from starting points close to the final parameter estimates. Poor convergence was noted when the number of iterations required ap- proached 20 or more and/or parameter estimates still changed substantially in later iterations. Coincidence of growth curves was tested using a general F-statistic of the form Foy —a.21k-1),pF,) = (OSS, — SS,)/2(k — 1I)V[SS,/DF,] where SS, = sum of squares for the single regression line for all populations combined, SS, = the sum of SS for regression lines fit to individual populations, k = number of populations compared, and Df, = degrees of freedom of SS, (Zar, 1996). If the F-statistic is not significant, this indicates that a single growth curve is adequate to de- scribe the populations compared. Tests of coincident growth curves (performed only for the normal Gompertz growth models) were made first for all sites combined The Veliger, Vol. 43, No. 1 Table 1 Age-class distribution (ind. 0.047 m~*) of Mytilus tros- sulus by site and year. B4-81 B4-82 MC-81 MC-82 SS-81 SS-82 Age No. No. No. No. No. No. 0) 268 128 12 428 ® 124 1 325 185 32 634 145 209 2 1345 644 354 130 554 216 3 402 667 1039 292 670 318 4 238 150 387 SD 345 515 5 233% = iytS5 163 245 154 201 6 175 41 ail 77 67 31 7 42 39 27 45 30 19 8 8 9 7 15 4 3 9 5 0) 2 9 1 10 1 l 1 5 1] 1 12 (0) 13 1 Total 3042 1919 2095 2457 2045 1636 and then, if the first F-statistic was significant, between pairs of sites. RESULTS The age-frequency distributions for mussels from the study sites indicated a dominance of young mussels (Ta- ble 1). Age class O mussels did not occur in large num- bers. The minimum count for this age class was 12 mus- sels 0.047 m~? in 1981, and the maximum was 428 mus- sels 0.047 m~? in 1982, all at the MC site. Mussels in age classes 1-3 were the most abundant, and the first six age classes (age 0 to 5) generally comprised over 90% of the individuals present at all sites. Mussels older than 7 years of age were relatively uncommon. It was only at the MC site that mussels older than 10 years occurred. The min- imum length observed was for an age 0 mussel of length 0.49 mm at the B4 site, and the maximum was 58.00 mm for an 8-year-old mussel at SS. Assessment of shell growth by age class indicates that the highest percentage of new shell growth occurs within the younger age classes with only a small percentage of new growth added after age 6 (Tables 2, 3, and 4). The new shell growth of mussels can represent over 50% of mean shell length for age class 1 mussels. Shell growth does not occur during winter months, from December to February, and remains minimal through March (Figure 2). The percentage of new shell growth per month in- creases greatly in May, and maximum growth rates, ap- parent as the steepest portions of the lines in the plots of % new shell growth, occur from June to August. The normal and seasonalized Gompertz growth models revealed some differences between sites (Figure 3). Con- vergence of the normal Gompertz models was excellent, A. Blanchard & H. M. Feder, 2000 Page 37 Table 2 Mean length and standard error (SE) of Mytilus trossulus in each age class by site and year. N/A = not available due to either no mussels or a single mussel in an age class. B4-81 B4-82 MC-81 Age Mean SE Mean SE Mean SE 0) 2.54 0.06 2.01 0.09 2.90 0.18 1 6.79 0.21 535 0.26 8.45 0.63 2 14.89 0.17 11.87 0.23 14.49 0.30 3 21.79 0.35 24.06 0.28 18.89 0.20 4 28.85 0.45 30.76 0.51 28.80 0.34 5 35.10 0.36 33.78 0.72 34.78 0.39 6 38.15 0.37 37.28 0.65 37.85 0.56 7 38.24 0.64 39.01 0.67 40.49 0.77 8 40.13 2.39 37.70 1.20 40.59 1.25 9 39.44 1.00 N/A N/A 46.67 DDS 10 44.82 N/A 35.12 N/A 48.75 N/A 11 12 — Ww MC-82 SS-81 SS-82 Mean SE Mean SE Mean SE 2.33 0.04 2.91 0.16 2223 0.13 3.20 0.07 7.16 0.29 5.63 0.17 9.29 0.32 13.81 0.30 12.18 0.38 18.80 0.29 23.85 0.30 21.84 0.43 24.16 0.20 26.95 0.46 29.48 0.28 31.84 0.31 Shia 0.69 35.00 0.36 36.73 0.52 36.29 1.01 38.57 0.72 38.90 0.70 40.83 1.05 40.35 1.26 40.59 3, 45.04 4.58 43.23 2.05 43.54 1.09 46.81 N/A 40.55 1.83 40.38 N/A N/A N/A 37.50 N/A but convergence of the seasonalized models was poorer, particularly for the SS site. Tests for coincidence of growth curves between sites were all highly significant (p < 0.0001) indicating that a curve for each site was necessary to estimate the normal growth models. Com- parison of the 95% confidence intervals of parameter es- timates between each site (Table 5) indicated that model estimates for the MC site were significantly different than the B4 and SS sites (observed as no overlap of the 95% confidence intervals), while the values of the B4 and SS sites were not different (i.e., overlap of confidence inter- vals for each parameter). A poor fit of the growth curves for age O mussels (Figure 3) is apparent as the mean lengths of age class 0 mussels from January to April (age < 0.4) was greater than the values estimated in the growth curves. Since movement into the next age class is marked by new shell growth, first visible in the spring, the age O mussels occurring in the first of the year (Jan- uary through April) are those approaching the period of new shell growth and their movement into the next age class. The seasonalized Gompertz model provides an esti- mate of two additional parameters, C and t,, useful for assessing growth patterns. The amplitude of the seasonal component estimated by the parameter C, here fixed at one based on preliminary analyses, indicates high ampli- tude seasonal growth patterns and suggests that no growth occurs during the period estimated by t,, (t, = t, + 0.5). Table 3 Mean (mm) new shell growth and standard error (SE) of Mytilus trossulus in each age class by year and by station. N/A = not available due to either no mussels or a single mussel in an age class. B4-81 B4-82 MC-81 Age Mean SE Mean SE Mean SE 1 3.94 0.17 332 0.20 SZ 0.47 2 6.32 0.12 4.28 0.17 6.18 0.24 3 6.09 0.22 4.93 0.19 4.82 0.14 4 4.32 0.23 BI 0.27 4.08 0.17 S) 3.08 0.16 1.38 0.27 2.00 0.16 6 1.98 0.13 0.97 0.16 1.18 0.15 7 1.08 0.22 0.64 0.11 1.08 0.27 8 0.91 0.29 0.29 0.13 1.55 0.40 9 0.19 0.13 N/A N/A 12 0.34 10 0.00 N/A 0.12 N/A 0.87 N/A 11 12 MC-82 SS-81 SS-82 Mean SE Mean SE Mean SE 1.87 0.06 4.25 0.23 3.66 0.15 2.34 0.23 5.70 0.20 4.71 0.29 2.26 0.17 S78} 0.15 2.96 0.20 Da 0.10 3.58 0.19 3.45 0.11 2.32 0.14 DIDS 0.18 2.32 0.15 1.45 0.14 1.41 0.18 1.28 0.25 1.23 0.20 0.93 0.16 1.10 0.25 0.58 0.14 0.66 0.14 0.50 0.31 0.51 0.11 0.59 N/A 0.41 0.15 0.16 N/A N/A N/A 0.20 N/A Page 38 The Veliger, Vol. 43, No. 1 Table 4 Percent new shell growth (of total length) of Mytilus tros- sulus in each age class by year and by station. N/A = not available due to either no mussels or a single mussel in an age class. B4-81 B4-82 MC-81 MC-82 SS-81 SS-82 Age %o %o % % % % 1 57.95 62.09 67.74 58.54 59.30 65.00 2 42.48 36.07 42.61 25.17 41.31 38.70 3 27.94 20.48 25.52 12.04 24.01 13.54 4 14.98 10.44 14.16 11.45 13.28 11.71 5 8.77 4.10 5.76 7.30 7.22 6.64 6 5.20 2.60 3.11 3.94 3.88 3.31 7 2.82 1.64 2.67 3.17 2.27 2/3. 8 2.26 0.78 3.82 1.44 1.47 1.16 9 0.49 N/A 2.58 1.17 1.26 10 0.00 0.34 1.78 1.01 11 0.40 12 N/A 13 0.53 The t, coefficients indicate that 49% to 53% of the year elapses before growth rates reach maximum values cor- responding to the month of June and thus, t, corresponds to January. DISCUSSION Mytilus trossulus in Port Valdez demonstrates shell growth patterns comparable to growth of Mytilus from Greenland (e.g., Theisen, 1973). Theisen (1973) consid- ered mussels in Greenland to be M. edulis, but based on current knowledge, those mussels could be either M. ed- ulis or M. trossulus (Gosling, 1992). Relative to the growth of mussels from the intertidal samples reported by Theisen (1973), mean shell lengths of 3 to 4 mm are recorded for mussels of age class 1 (as determined by applying our aging technique to his growth check data), 30 to 36 mm for mussels for 6-year-old mussels, and 42 to 52 mm for 9-year-old mussels. Lengths of the first few age classes of the present study (e.g., age O—3 mussels; Table 1) are greater than those from the Theisen (1973) study, but mean lengths of mussels after the sixth year are similar. Intertidal mussels of Theisen (1973) demon- strated life spans similar to those observed in this study (up to 12—13 years), and younger mussels were dominant in the Greenland samples as well, with mussels >7—8 years of age relatively uncommon. The range of maxi- mum length of mussels estimated by the Gompertz model from sites in Port Valdez (41.0 to 44.3 mm) is less than those estimated by Theisen (1973) (54.9 to 62.5 mm). Unlike the present study where the Gompertz model fit all age classes relatively well, Theisen concluded that the von Bertalanffy growth equation fit his data better for older mussels, while the Gompertz equation was better for the younger mussels. The differences in model fitting and the estimated maximum lengths may be due to the varying methodologies (the Ford-Walford plot method vs. nonlinear regression) or possibly to genetic differences in growth between M. trossulus and M. edulis. Decreased temperature and salinity are known to result in reduced growth in mussels (Seed & Suchanek, 1992). Theisen (1973) reported reduced shell growth of mussels from Greenland, as compared to populations at lower lat- itudes, and related reduced shell growth to low tempera- tures within his study area. Tedengren & Kautsky (1986) concluded that the reduced growth of Baltic Sea mussels (now considered to be M. trossulus: Gosling, 1992), com- pared to mussels from the North Sea, resulted from phys- iological adjustments to low salinity (from 4.5 to 10%— similar to conditions in Port Valdez in summer). Thus, shell lengths of mussels from Port Valdez may be reflec- tive of seasonal environmental stress. Natural sources of stress at the study sites in Port Valdez include high sed- iment loads (ranging up to 100 mg L~'; Sharma & Bur- bank, 1973) and decreased salinity (approaching 0%oc; Blanchard & Feder, 1997) in summer, and freezing air temperature in winter, as well as year-round low water temperatures. However, to fully understand the effects of decreased temperature and salinity on shell growth of M. trossulus in Port Valdez and along the Pacific coast, ex- perimental studies similar to those performed for M. ed- ulis are necessary (e.g., see review by Seed & Suchanek, 1992). The seasonal growth pattern in mussels from Port Val- dez reflects the subarctic climate at this fjord. In summer, the length of daylight approaches a maximum of 20 hours while in winter, from mid November to early February, there are less than 7 hours (Alexander & Chapman, 1980). Phytoplankton abundance increases in early spring, as daylight increases, reaching maximum levels in May and June (Alexander & Chapman, 1980). A minor phytoplankton bloom occurs in fall, but low abundance levels are reached by November and continue through the winter. Consequently, shell growth in Mytilus is minimal in spring, increases rapidly in mid-summer once gamete maturation is completed and spawning initiated, and ceas- es in winter (Feder & Keiser, 1980; Blanchard & Feder, 1997). This seasonal growth pattern is indicated by the plots of % new shell growth (Figure 2) and the season- alized Gompertz growth models (Figure 3). Due to the response of shell growth to varying condi- tions in field studies, it may be impossible to discriminate any sublethal effects of contaminants from natural envi- ronmental variations. Studies attempting to assess effects of contaminants on Mytilus generally rely on assessment of other aspects of mussel biology. Blanchard & Feder (1997) investigated the reproductive and nutritive storage tissue cycles for M. trossulus in Port Valdez including the sites of the present study. They found that mussels at the MC site demonstrated significantly less reproductive tis- A. Blanchard & H. M. Feder, 2000 Page 39 Gametic and Nutr. VW. 7) © s a a = PS = s 2 3 ¢ © o s ® £ © to) WW Gametic and Nutr. Tissues Figure 2 Bar charts of proportions of gametic tissue overlain by proportions of nutritive tissues (data from Blanchard & Feder, 1997) and percent new shell growth for Mytilus trossulus from Port Valdez in 1981 and 1982. Filled bars indicate the first occurrence of appreciable amounts of reproductive tissues in spawning condition. Page 40 The Veliger, Vol. 43, No. 1 Table 5 Parameter values and 95% confidence intervals for Gompertz growth models. L., = maximum length, k = growth coefficient, t) = length at time = O, and t, = fraction of year before maximum growth rate. ASE = asymptotic standard error. N/A = not available. Parameter Site Parameter estimate B4 ex 40.98 k 0.668 (5 2.651 Seasonal Model ee 41.28 k 0.664 ty 2.694 t, 0.491 MC ee 44.30 k 0.511 tG 3.131 Seasonal Model [ez 43.74 k 0.525 ty 3.079 t. 0.533 SS L. 41.55 k 0.608 to 2.731 Seasonal Model* ee 42.26 k 0.605 te 2.788 t, 0.508 95% Confidence intervals Lower Upper ASE bound bound 0.404 40.18 41.77 0.015 0.639 0.696 0.019 2.614 2.688 0.395 40.51 42.06 0.014 0.637 0.692 0.019 2.656 DBD 0.017 0.459 0.524 0.558 43.21 45.40 0.011 0.490 0.532 0.030 3.072 3.190 0.525 42.71 44.77 0.011 0.504 0.546 0.029 3.023 3.135 0.017 0.500 0.566 0.693 40.19 42.91 0.020 0.569 0.647 0.034 2.664 2.797 N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A * Convergence of this growth curve was particularly poor so standard errors and confidence intervals are not presented. The parameter estimates should be considered as approximate values. sue than did the B4 and SS sites and suggested that mus- sels at the MC site were negatively influenced by the colder, silt-laden waters of Mineral Creek. They further concluded that effects of hydrocarbon contamination, de- rived from small spills and industrial activities in the ter- minal area, were not apparent in the reproductive cycle of mussels from Port Valdez. Similarly, the growth curves presented here revealed no differences that could be clearly related to the proximity of mussels at B4, within the terminal area, to hydrocarbon contamination. The overlap of the 95% confidence intervals of curve param- eters for the B4 and SS sites indicates that although the curves for the two populations are not coincident (.e., the F-statistic was significant), the curves are still very similar. The significant differences in parameter estimates between the curve for the MC sites and the curves for B4 and SS sites, are suggestive of environmental differ- ences between the MC site on the northern shore and the B4 and SS sites on the southern shore of Port Valdez. Additional conclusions can be drawn from the age-fre- quency distributions. Recruitment by young-of-the-year mussels was originally thought to occur by secondary set- tlement into the mussel beds, but recent evidence indi- cates that these juveniles may also settle directly into the beds (McGrath et al., 1988; Seed & Suchanek, 1992). Feder & Keiser (1980) observed that newly settling mus- sels (early plantigrade stage, < 0.4 mm; Seed & Sucha- nek, 1992) in Port Valdez generally settled on filamentous algae and then moved into the mussel beds. All age 0 mussels in the present study were larger than the early plantigrade stage suggesting initial settlement outside of mussel beds. Additionally, the typically low numbers of age O mussels and large numbers of age classes 1 and 2 mussels support the secondary settlement hypothesis of Feder & Keiser (1980). While direct recruitment into the mussel beds was not observed during the 1981-1982 study, an investigation of the abundance of mussels in Port Valdez from 1989-1992 (Feder & Blanchard, 1993) indicated that large numbers of plantigrade mussels can sporadically settle directly into mussel beds (up to 2200 individuals in 0.016 m? replicates in 1990: unpublished data). Also, the age-frequency distributions indicated that the mussels in Port Valdez are generally short-lived with mussel beds dominated by mussels 5-years-old or less, and older mussels progressively more uncommon. The findings of this study reveal a well-defined sea- sonal shell-growth pattern for M. trossulus in Port Valdez. The growth patterns of mussels reflect the extreme en- vironmental conditions encountered within the intertidal region of Port Valdez. The seasonalized Gompertz growth A. Blanchard & H. M. Feder, 2000 Page 41 h4 G5 Gompertz Growth Model Bert 50 “™. Seasonal Gompertz Model e Mean Length . r Mean Length (mm) lass: Gompertz Growth Model Mineral Creek 50 “Seasonal Gompertz Model e Mean Length Mean Length (mm) iss Gompertz Growth Model Sawmill Spit 60 ™.~. Seasonal Gompertz Model e Mean Length ° Mean Length (mm) Age Figure 3 Gompertz growth curves and mean shell lengths for Mytilus trossulus from the (a) Berth 4, (b) Mineral Creek, and (c) Sawmill Spit sites in 1981-1982 from Port Valdez. Age is calculated as the sum of the observed age (by counting growth rings) and the fraction of the year in which sampling occurred. The tick marks denote fractions of a year. Page 42 curves indicate a strong seasonal component to shell growth with maximum shell growth rates during and/or after June; most growth occurs during mid summer after spawning commences. Differences in growth curves be- tween the MC site and the B4 and SS sites are reflective of environmental differences between sites on opposite sides of the bay. Effects on shell growth due to hydro- carbon contamination were not apparent. Age-frequency distributions suggest secondary settlement of juveniles into mussel beds as the primary method for recruitment. Acknowledgments. We thank the many technicians who col- lected and measured mussels. Funding for this study was made available by Alyeska Pipeline Service Company. We would like to thank Dr. B. Roth, Dr. R. Seed, and an anonymous reviewer for their constructive comments. LITERATURE CITED ALEXANDER, U. & T. CHAPMAN. 1980. Phytotoxicity. Pp. 125— 142 in J. M. Colonell & H. K. Stockholm (eds.), Port Valdez, Alaska. Environmental Studies, 1976-1979. Institute of Ma- rine Science Occasional Publication No. 5, University of Alaska: Fairbanks, Alaska. BLANCHARD, A. & H. M. FEDER. 1997. Reproductive timing and nutritional storage cycles of Mytilus trossulus Gould, 1850, in Port Valdez, Alaska, site of a marine oil terminal. The Veliger 40:121—130. COLONELL, J. M. 1980. Physical oceanography. Pp. 9-36 in J. M. Colonell & H. Stockholm (eds.), Port Valdez, Alaska: En- vironmental Studies 1976-1979. Institute of Marine Science: Fairbanks, Alaska. FEDER, H. M. & A. BLANCHARD. 1993. A long-term intertidal monitoring program for Port Valdez, Alaska 1991-1992. Fi- nal report to Alyeska Pipeline Service Company, Institute of Marine Science, University of Alaska, Fairbanks. Pp. 275. Available from Environment & Natural Resources Institute, University of Alaska, 7707 A St., Anchorage, Alaska 99501. Feber, H. M. & G. E. KEISER. 1980. Intertidal biology. Pp. 143-— 224 in J. M. Colonell & H. K. Stockholm (eds.), Port Valdez, Alaska. Environmental Studies, 1976-1979. Institute of Ma- The Veliger, Vol. 43, No. 1 rine Science Occasional Publication No. 5, University of Alaska: Fairbanks, Alaska. GOSLING, E. M. 1992. Systematics and geographic distribution of Mytilus. Pp. 1-20 in E. M. Gosling (ed.), The Mussel My- tilus: Ecology, Physiology, Genetics and Culture. Elsevier Science Publishers: Amsterdam. McDOona_Lp, J. H., R. SEED & R. K. KOEHN. 1991. Allozymes and morphometric characters of three species of Mytilus in the Northern and Southern Hemispheres. Marine Biology 111:323-333. McGratH, D. P. A. KING, & E. M. GOSLING. 1988. Evidence for the direct settlement of Mytilus edulis larvae onto adult mus- sel beds. Marine Ecology Progress Series 47:103—106. RATKoSWKy, D. A. 1990. Handbook of Nonlinear Regression. Marcel Dekker, Inc.: New York. 241 pp. SEED, R. & T. H. SUCHANEK. 1992. Population and community ecology of Mytilus. Pp. 87-169 in E. M. Gosling (ed.), The Mussel Mytilus: Ecology, Physiology, Genetics and Culture. Elsevier Science Publishers: Amsterdam. SHARMA, G. D. & D. C. BURBANK. 1973. Geological oceanog- raphy. Pp. 223—250 in D. W. Hood, W. E. Shiels & E. J. Kelley (eds.), Environmental Studies of Port Valdez. Insti- tute of Marine Science Occasional Publication No. 3, Uni- versity of Alaska: Fairbanks, Alaska. SHaAw, D. G. & M. J. HAMEEDI. (eds.). 1988. Environmental Stud- ies in Port Valdez, Alaska. Springer-Verlag: New York. 423 Pp. SPARRE, P., E. URSIN. & S. C. VENEMA. 1989. Introduction to tropical fish stock assessment. FAO Fisheries Technical Pa- pers 306/1. P. 337. Spss INc. 1997. SYSTAT 7.0. SPSS Inc.: Chicago, Illinois. STROMGREN, T., M. V. NIELSEN, & K. UELAND. 1986. Short-term effect of microencapsulated hydrocarbons on shell growth of Mytilus edulis. Marine Biology 91:33-39. TEDENGREN, M. & N. KAutTsky. 1986. Comparative study of the physiology and its probable effect on size in blue mussels (Mytilus edulis L.) from the North Sea and the northern Baltic proper. Ophelia 25:147—155. THEISEN, B. F 1973. The growth of Mytilus edulis L. (Bivalvia) from Disko and Thule district, Greenland. Ophelia 12:59— Ws Zar, J. H. 1996. Biostatistical Analysis. 3rd ed. Prentice Hall: Upper Saddle River, New Jersey. 662 pp. The Veliger 43(1):43—SO (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Two New Neogene Species and the Evolution of Labral Teeth in Concholepas Lamarck, 1801 (Neogastropoda: Muricoidea) THOMAS J. DEVRIES Box 13061, Burton, Washington 98013, USA Abstract. Two new species of Concholepas Lamarck, 1801, are described from southern Peru, Concholepas chiro- tensis sp. nov. (middle Miocene) and Concholepas camerata sp. nov. (late Pliocene). Both species inhabited high-energy nearshore environments, as have most species of Concholepas. The evolution of the genus was most rapid during the late middle Miocene and the late Pliocene, times when the molluscan fauna of the Peruvian Faunal Province experienced mass extinction. A case is made that the late Pliocene development of labral teeth in Concholepas was related to changes in prey precipitated by global cooling. INTRODUCTION The fossil record of the muricid gastropod genus Con- cholepas Lamarck, 1801, was reviewed recently by DeVries (1995). The known geographic ranges of two Chilean Pliocene species, C. nodosa Moricke, 1896, and C. kieneri Hupé, 1854, were extended northward to the Pisco and Sacaco Basins of south-central Peru, and the temporal range of the latter species back to the late Mio- cene. A new early to middle Miocene species from Peru, C. unguis DeVries, 1995, was also described. Samples recently collected between Pisco and Camana (Figure 1) have yielded two new species of Concholepas. C. chi- rotensis sp. nov. represents a middle Miocene transition from C. unguis to C. kieneri. C. camerata sp. nov., from upper Pliocene strata, exhibits traits of both C. kieneri and the extant C. concholepas (Bruguiére, 1789), includ- ing the two labral teeth of the latter species. GEOLOGY Upper Miocene, Pliocene, and Pleistocene marine depos- its are widely exposed in the Pisco and Sacaco Basins (Figure 1; Muizon & DeVries, 1985; DeVries, 1998). To the south, lower to middle Miocene bioclastic sandstones of the Camana Formation surround Camana (Pecho & Morales, 1969; Ibaraki, 1992). Deposits that unconform- ably overlie the Camana Formation were assigned to the upper Miocene Pisco Formation, but the assemblage of mollusks listed by Pecho & Morales (1969) suggests an early to middle Miocene age. Between the Sacaco Basin and Camana, scattered outcrops of bioclastic sandstone and conglomerate occur along the coast (Olchauski, 1980). Mollusks from these deposits reported by Beaudet et al. (1976) and others found by this author (Chorus grandis [Philippi, 1887] and Acanthina unicornis [Bru- guiére, 1789]) indicate a late Pliocene to early Pleistocene age. Abbreviations used for localities or specimens are as follows: USNM, Department of Paleobiology, United States Na- tional Museum of Natural History, Washington, D.C., USA; L, anterior-posterior Length; W, width at widest point, perpendicular to length; THK, dorso-ventral thick- ness, measured at widest point. Material is described with a collections number, local- ity/sample number, and dimensions (L, W, THK). Mea- surements enclosed by parentheses indicate sizes for bro- ken specimens. Locality descriptions are given in the ap- pendix. SYSTEMATIC PALEONTOLOGY Family MurIcIDAE Rafinesque, 1815 Subfamily RAPANINAE Gray, 1853 Genus Concholepas Lamarck, 1801 Concholepas Lamarck, 1801: 69. Type species (monotypy): Concholepas peruviana La- marck, 1801 (= Buccinum concholepas Bruguiére, 1789). Concholepas camerata DeVries, sp. nov. (Figures 2, 5, 6, 9-11, 14) Diagnosis: Adults with highly arched body whorl, quad- rate aperture; two prominent labral teeth on outer lip. Description: Shell to 100 mm long, narrow, ovate. Body whorl extending full length of adult shell. Spire sub- merged. Protoconch not preserved. Body whorl of adults with arched to tabular sutural platform, steeply rounded shoulder, and deeply vaulted mid-section resulting from diminished whorl expansion rate in last quarter to half turn of body whorl. Periphery at midpoint of whorl; an- terior longitudinal profile steeply descending to siphonal The Veliger, Vol. 43, No. 1 75°W — PANAMERICAN C2] Pisco Basin 0 100 km _—_— | SCALE Figure 1 Extent of the Pisco Basin in south-central Peru. The smaller Sa- caco Basin extends from a few kilometers north of Sacaco to Yauca. Lower and middle Miocene marine sediments outcrop around Camana. notch; no anterior constriction. Juveniles variably inflat- ed. Fasciolar ridge strong; pseudumbilical area dorso-ven- trally broad, planar; extending three-quarters length of shell. Aperture quadrate; inner lip slightly reflected out- ward; columella broad, planar, posterior interior edge bladelike. Posterior end of outer lip inflected at shoulder. Outer lip coarsely crenulate. Spiral sculpture consisting of eight to nine low rounded spiral cords, with one or two secondary cords in most interspaces. Primary spiral cords with irregularly spaced low nodes, sometimes frilled on early whorls where crossed by growth lines. Anteriormost two interspaces produced and thickened from within outer lip to form two labral teeth. Discussion: The sculptural pattern of C. camerata is un- like that shown by C. kieneri (compare Figures 2 and 3) but resembles that of C. concholepas: rounded primary spiral cords that are sometimes lamellose; few secondary cords; and two strong labral teeth. The aperture of most specimens of C. camerata, however, is more like that of C. kieneri (compare Figures 6, 7, and 8): quadrate, not oval; and with a columella whose posterior interior edge is bladelike, not rolled under and thickened, as it is in C. concholepas; and an inner lip that is nearly upright, not strongly reflected. The pattern of whorl expansion for Concholepas ca- merata is intermediate between that seen in C. kieneri and C. concholepas (compare Figures 11, 12, and 13). In C. kieneri, the expansion rate regularly changes, produc- ing a boxlike transverse profile. In C. camerata, more rapid juvenile expansion rates are succeeded by lower adult rates, producing a deeply arched transverse profile. In C. concholepas, the juvenile expansion rate persists through adult stages, producing a flatter, uniformly con- vex transverse profile. A juvenile specimen from locality 95DV 812 (USNM 447126, Figures 9 and 14) exhibits the broad curvature and reflected inner lip of C. concholepas, but still retains the shoulder inflection and sharp inner edge at the pos- terior end of the columella seen in C. camerata. Other juvenile specimens from locality 95DV 812 (e.g., USNM 447128, Figures 5 and 10) show the full suite of char- acteristics diagnostic of C. camerata. Type locality: Locality 95DV 812, south side of Que- brada Huacllaco, roadcut along Panamerican Highway (Figure 21). Specimens of C. camerata come from small inliers of sandy coquinas that overlie basement crystalline Explanation of Figures 2—20 Figures 2, 5, 6, 9-11, 14. Concholepas camerata DeVries, sp. nov. All from sample 9SDV 812-1, Quebrada Huacllaco, late Pliocene. 2. USNM 447129, holotype, abapertural view, 0.7. 5. USNM 447128, paratype, abapertural view, 1.5. 6. USNM 447129, apertural view, X0.7. 9. USNM 447126, paratype, aba- pertural view, <1.0. 10. USNM 447128, abapertural view, 1.0. 11. USNM 447129, view from posterior showing transverse pro- file, 0.75. 14. USNM 447126, apertural view, 1.0. Figures 3, 7, 12. Concholepas kieneri Hupé, 1854. USNM 447088, sample 86DV 360-1, South Sacaco, early Pliocene. See DeVries (1995) for more locality data. 3. Abapertural view, «0.48. 7. Apertural view, X0.52. 12. View from posterior show- ing transverse profile, *0.52. Figure 4. Concholepas nodosa Moricke, 1896. USNM 447122, sample 95DV 809-1, Yauca, early Pliocene, abapertural view, «1.88. Figures 8, 13. Concholepas concholepas (Bruguiére, 1789). Cen- tral Peru, Recent. 8. Apertural view, <0.5. 13. View from pos- terior showing transverse profile, 0.50. Figures 15-19. Concholepas chirotensis DeVries, sp. nov. Ca- mana, middle Miocene. 15. USNM 447120, holotype, sample 95DV 816-1, abapertural view, <1.0. 16. USNM 447120, aper- tural view, X1.0. 17. USNM 447123, sample 9SDV 815-1, aba- pertural view, broken specimen, X1.0. 18. USNM 447121, par- atype, sample 95DV 816-1, abapertural view, 1.0. 19. USNM 447121, apertural view, 1.06. Figure 20. Concholepas unguis DeVries, 1995. USNM 447096, holotype, sample 87DV 579-2, near Fundo Santa Rosa, abaper- tural view, X1.0. See DeVries (1995) for more locality data. Page 45 T. J. DeVries, 2000 The Veliger, Vol. 43, No. 1 me ne mS a) HALA “™ OF —— HIGHWAY oS 00 5 409 15°55'S WY w ne) ron) a fo 3 Ce Figure 21 Type locality of Concholepas camerata DeVries, sp. nov. rock at elevations of about 200 to 250 meters above sea level on a narrow wavecut platform. In addition to specimens from the type locality, spec- imens of C. camerata are also known from upper Plio- cene beds in the Sacaco Basin (Localities 96DV 911 and 96DV 923), where they overlie balanid coquinas contain- ing specimens of C. kieneri and underlie similar coquinas containing specimens of C. concholepas. Etymology: From the Latin word ‘‘camera,” “‘vaulted chamber.”’ meaning Material: (Sample 95DV 812-1 for all specimens) Ho- lotype USNM 447129, L 79.5 mm, W 56.7 mm, THK 37.1 mm; paratype USNM 447125, L 97.8 mm, W (63) mm, THK (49.4) mm; paratype USNM 447126, L 40.2 mm, W 29.9 mm, THK 12.9 mm; paratype USNM 447127, L 36.0 mm, W (19) mm, THK 14.8 mm; USNM 447128, L 15.5 mm, W 11.4 mm, THK 5.6 mm. Occurrence: Upper Pliocene, southern Peru. Concholepas chirotensis DeVries, sp. nov. (Figures 15—19) Diagnosis: Shell small, evenly inflated; low spire; pseud- umbilical area 60 percent of shell length; surface sculp- ture of numerous spiral cords, including five to seven nodular primary spiral cords. Description: Shell small, to 35 mm; ovate, with a large aperture (95 percent of the shell length) and low spire. Spire consisting of two whorls; protoconch unknown. Body whorl without pronounced shoulder; whorls ap- pressed at suture; periphery posterior to midpoint of whorl; anterior quarter of body whorl sometimes weakly constricted. Fasciolar ridge strong, posterior edge with thin keel; pseudumbilical area usually as narrow as fas- ciolar ridge is wide, extending about 60 percent of shell length. Columella excavated, anterior end flattened, co- mea o/72°40'W SVs ge Figure 22 Type locality of Concholepas chirotensis DeVries, sp. nov. planar with aperture; posterior end twisted vertically be- neath ridge bordering pseudumbilical region. Inside of outer lip crenulate. Surface sculpture of about 50 spiral cords, including five to seven nodular primary cords, with three to seven secondary cords in interspaces. Numerous evenly spaced growth lines intersecting spiral cords to produce beaded texture. Discussion: Specimens of C. chirotensis have a pseud- umbilical area twice as long as that in specimens of the geologically older C. unguis and primary spiral cords that are more pronounced and nodular, rather than smooth (compare Figures 15, 17, 18, and 20). Compared with specimens of the geologically younger C. kieneri, those of C. chirotensis are much smaller, more uniformly in- flated, and have many more secondary spiral cords (com- pare Figures 3, 15, and 17). Type locality: Locality 95DV 816, Quebrada Chiroteo, 0.5 km east of juncture with Quebrada Mal Paso; roadcut on east side of the Panamerican Highway (Figure 22). At localities 95DV 815 and 95DV 816, roadcuts ex- pose an irregular granitic basement overlain by cross-bed- ded coarse-grained sandstones, lenses of gravel with specimens of C. chirotensis, laminae of heavy mineral sands, and, basally, rounded and angular cobbles of gran- ite with interspersed large valves of oysters. Two kilo- meters downhill along the Panamerican Highway, Ibaraki (1992) collected planktonic foraminifera from underlying T. J. DeVries, 2000 strata that yielded an age of late early Miocene (sample Pe-86-21). In addition to specimens from the type locality, internal molds of C. chirotensis are found in bioclastic sandstones overlying volcanic rocks in the northern Pisco Basin (lo- cality 97DV 935). Nearby outcrops of fossiliferous ma- rine sediments in the Filudo Depression have been as- signed a middle Miocene age (DeVries, 1998). Etymology: Named after Quebrada Chiroteo. Material: Holotype USNM 447120, 95DV 816-1, L 30.9 mm, W 21.0 mm, THK 14.3 mm; paratype USNM 447121, 9S5DV 816-1, L 24.3 mm, W 17.2 mm, THK 10.3 mm; USNM 447124, 95DV 816-1, six specimens ranging in length from 8.8 to 23.1 mm; USNM 447123, 95DV 815-1, L (38) mm, W 30.7 mm, THK (17) mm. Occurrence: Middle Miocene, southern Peru. DISCUSSION The pattern of evolution for Concholepas emerges more clearly with the addition of two species to the four al- ready known (Figure 23)—a pattern of gradual change, followed by two cycles of rapid (punctuated?) change and stasis. From the late early Miocene, by which time C. unguis had evolved from a probable non-Concholepas species, through the middle Miocene, the size of Con- cholepas shells doubled and the sculpture differentiated into primary and secondary cords (C. chirotensis), while coiling remained relatively tight. During the late middle Miocene, coincident with a major molluscan extinction in Peru and Chile, shell size tripled. Whorl expansion be- came alternatingly rapid and slow in each specimen, pro- ducing a dorso-ventrally flattened and quadrate transverse profile (C. kieneri). From the late Miocene through the early Pliocene, a time of relative faunal stasis in Peru, the proportions of C. kieneri remained unchanged. By the early Pliocene, however, a flattened form had evolved with a flaring aperture and a small number of lamellose primary spiral cords (C. nodosa; USNM 447122, Figure 4). By the late Pliocene, another time of mass extinction within the Peruvian molluscan fauna, both dorso-ventrally flattened species were extinct, replaced by populations with toothed shells, quadrate peristomes, and rapid rates of whorl expansion during juvenile stages (C. camerata). Toothed individuals with ovate peristomes, reflected inner and outer lips, and rapid rates of whorl expansion at all life stages (C. concholepas) appeared by the early Pleis- tocene. Four described genera of muricids in the Peruvian Fau- nal Province are distinguished by labral teeth (DeVries, 1997; DeVries & Vermeij, 1997). Unlike the single labral tooth of the three ocenebrine genera (Acanthina Fischer von Waldheim, 1807; Chorus Gray, 1847; Herminespina DeVries & Vermeij, 1997), which are formed by an in- folding and forward projection of the outer shell layer, Page 47 chirotensis camerata concholepas unguis Quatenary Late Pliocene Pliocene Late Miocene Middle Miocene Early Miocene Figure 23 Stratigraphic ranges and inferred phylogeny of Concholepas. The taxon ancestral to C. unguis remains unknown. An absence of specimens from upper Middle Miocene strata also leaves the na- ture of the transition from C. chirotensis DeVries, sp. nov. to C. kieneri in doubt. Ma = millions of years before present. the paired labral teeth of the rapanine Concholepas are thickened, tapered, blunt extensions of the two anterior- most interspaces. The teeth are used during feeding: to brace mytilids while opening them (M. Rabi, personal communication, 1995), to pry apart the integumental plates of barnacles, and to help push and crush (“‘bull- doze’’) small barnacles from rocky surfaces (Castilla et al., 1979). The prey favored by Concholepas varies with age and location, but includes barnacles (Balanus laevis Bruguiére, 1789; B. psittacus [Molina, 1782]; Chthalmus cirratus Darwin, 1854; C. scabrosus Darwin, 1854), my- tilids (Perumytilus purpuratus [Lamarck, 1819] Semimy- tilus algosus [Gould, 1850]; Brachidontes granulata [Hanley, 1843]); and the ascidian Pyura chilensis Molina, 1782 (Castilla et al., 1979; Dye, 1991; Moreno et al., 1993; DiSalvo & Carriker, 1994). The transition from untoothed to toothed populations of Concholepas is well preserved in a continuous 15-m section of alternating massive sandstones and bioclastic conglomerates on the southeastern flank of the Rio Acari Page 48 (Locality 96DV 923). Fossiliferous horizons in the lower 6.2 meters contain species (Anadara aff. A. chilensis [Phi- lippi, 1887]; Herminespina saskiae DeVries & Vermeij, 1997; Concholepas kieneri; C. nodosa) that indicate an early Pliocene age (Muizon & DeVries, 1985). Coquinas in the uppermost 3 meters contain species (Glycymeris ovata [Broderip, 1832]; Mulinia edulis [King, 1831]; Xanthochorus cassidiformis [Blainville, 1832]; Concho- lepas concholepas; Oliva peruviana Lamarck, 1811) that collectively indicate an early Pleistocene age (Muizon & DeVries, 1985). At 9 meters from the base of the section, cobble beds contain the highly arched toothed shells of Concholepas camerata. Specimens of C. kieneri are still found at 11 m, however, suggesting that populations of Concholepas without labral teeth persisted sympatrically for some tens of thousands of years after the toothed spe- cies evolved. Eighty percent of the marine mollusks from southern Peru and Chile, including all that might be considered ““tropical,’’ became extinct during the time (3—2 Ma) that toothed populations of Concholepas evolved (Herm, 1969; DeVries, unpublished data). This time was char- acterized by global cooling at middle and high latitudes (Dowsett et al., 1996). As may have been the case else- where (Stanley, 1986; Petuch, 1995), decreasing sea sur- face temperatures in the southeastern Pacific Ocean prob- ably precipitated the mass extinction of Peruvian mol- lusks. Concomitant with the mass extinction and evolution of labral teeth in Concholepas, the relative abundance of balanid species at Locality 96DV 923 changed. Lower Pliocene strata contain the scuta of Balanus laevis and B. tintinnabulum concinnus Darwin, 1854. Upper Pliocene strata, seemingly no different in lithology, location, or paleogeographic setting, additionally contain numerous scuta of B. psittacus. Unfortunately, no mytilids were found in the Rio Acari deposits, so it cannot be deter- mined if other prey species of Concholepas varied in abundance during the same time. It is instructive to examine the regions from which modern prey species of Concholepas were probably re- cruited during the late Pliocene. Many of the attached rocky intertidal species of the modern Peruvian Faunal Province upon which Concholepas preys range well into the cold-water Magellanic Province of southern South America. Balanus psittacus ranges from central Peru to the Straits of Magellan (Pilsbry, 1916). Perumytilus. pur- puratus ranges from Ecuador to the Straits of Magellan and north to Santa Cruz, Argentina (Soot-Ryen, 1959). Brachidontes granulata, Chthalmus scabrosus, and Bal- anus tintinnabulum concinnus all range as far south as southern Chile (Pilsbry, 1916; Soot-Ryen, 1959). Semi- mytilus algosus ranges from northern Peru only as far south as south-central Chile (Marincovich, 1973), but a southern origin is suggested for this mytilid by its steep decline in numbers when warm water spread along the The Veliger, Vol. 43, No. 1 coast of Peru during the 1982—1983 El] Nifio event (Arntz & Tarazona, 1990). In summary, the modern rocky inter- tidal attached fauna of Peru has a decidedly cold-water cast, more so than the vagrant rocky intertidal fauna or sandy bottom fauna, which have greater proportions of endemic species and immigrant species from warm wa- ters of the Panamic Province. The following hypothesis is proposed to explain the evolution of the labral teeth in Concholepas in a context of the ecological disruption associated with the late Pli- ocene mass extinction and in light of the modern feeding behavior of C. concholepas. Prior to the late Pliocene, less structurally sturdy species of Concholepas inhabited the subtidal and lower intertidal zone from southern Peru to central Chile. Individuals of C. kieneri and C. nodosa probably fed upon a variety of endemic small barnacles and ascidians, as do adults and newly recruited juveniles of C. concholepas today (Castilla et al., 1979; Moreno et al., 1993). During the late Pliocene, coastal waters cooled. At the same time, much of the coastal plain of Peru and Chile ceased subsiding and began to rise, producing a coastline that was straighter, rockier, and exposed to high- er-energy waves (DeVries, 1985, 1986, 1988). As these changes took place, larvae of mytilids and barnacles swept northward by the Peru-Chile Current from the long-standing rocky shores of southern Chile would have found acceptable substrates for colonization in suitably cool waters. Juvenile individuals of Concholepas may have gradually begun feeding upon newly introduced cold-water mytilids and balanids found higher in the in- tertidal zone, as juveniles do today (Dye, 1991; Moreno et al., 1993). Selection might then have favored arched and corrugated shells with labral teeth capable of dis- lodging and gripping prey in high-energy intertidal set- tings. The appearance of labral teeth appears not to be an improbable event in muricids (DeVries & Vermeij, 1997). A single specimen of the early Pliocene Chorus grandis (USNM 447074; DeVries, 1997; pl. 2, fig. 6a, b), for example, developed a second, incompletely formed labral tooth anterior to the single fully developed tooth that characterizes all species of the genus. This evolutionary scenario, in which labral teeth de- veloped in response to changes in prey induced by cli- mate change, could apply equally well to populations of C. kieneri from southern Peru or northern Chile. It is conceivable that a Pliocene population of C. kieneri in southern Chile might have first evolved labral teeth. Lar- vae of C. camerata might then have been carried north in the Peru Coastal Current together with larvae of cold- water balanids and mytilids. There is no evidence, how- ever, for any Miocene or Pliocene species of Concholepas in southern Chile (Hupé, 1854; Philippi, 1887; Herm, 1969; Watters & Fleming, 1972; Tavera, 1979; DeVries et al., 1984; Covacevich & Frassinetti, 1990; Frassinetti, unpublished data on collections from upper Miocene de- posits on Isla Mocha, south-central Chile). The occur- T. J. DeVries, 2000 rence of Holocene examples of C. concholepas in South Africa (Kensley, 1985) circumstantially suggests that its range has become more extensive at high latitudes fol- lowing its evolution at lower latitudes. The hypothesis for the evolution of labral teeth in Con- cholepas can be tested. A careful search for fossil mytil- ids from rocky intertidal deposits in Peru might reveal their biogeographic history. Quantitative studies of fossil barnacle diversity, biogeography, and predation in Peru and Chile might establish the role of their distribution in the evolution of Concholepas. Finally, experimental stud- ies on modern C. concholepas could address the adaptive significance of the labral teeth. Acknowledgments. I would like to thank V. Alleman of the Fa- cultad de Ciencias Biologia, Universidad Ricardo Palma (URP), Lima Peru, for providing logistical assistance and for her collab- oration in the field; M. Rabi, a consultant for the Instituto del Mar del Peru, for his insight into the behavior of Concholepas; the Facultad de Ciencias de Computadores (URP) for the use of university computer facilities; and Sr. and Sra. Carlos Martin de Buey for their hospitality. G. Vermeij and an anonymous review- er provided helpful suggestions in their critique of this manu- script. This research was funded by the author. LITERATURE CITED ARNTZ, W. E. & J. TARAZONA. 1990. Effects of El Nino 1982— 1983 on benthos, fish, and fisheries off the South American Pacific coast. Pp. 323-360 in P. W. Glynn (ed.), Global Eco- logical Consequences of the 1982—1983 El Nino-Southern Oscillation. Elsevier: Amsterdam. BEAUDET, G., D. HERM, R. LAHARIE & R. PASKOFF. 1976. Sur l’existence du Pliocene le long de la céte du sud du Pérou. Comptes rendus, sommaires de la Société Géologique Fran- cais 1976(1):12-13. CASTILLA, J. C., CH. GuIsADO & J. CANCcINO. 1979. Aspectos ecoldgicos y conductales relacionados con la alimentacion de Concholepas concholepas (Mollusca: Gastropoda: Muri- cidae). Biologia Pesquera Chile 12:99—114. COovACEVICH, V. & D. FRASSINETTI. 1990. La fauna de Lo Abarca: Hito biochronoestratigrafico y paleoclimatico en el Terciario superior marino de Chile central. 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Oligocene deposition and Cenozoic se- Page 49 quence boundaries in the Pisco Basin (Peru). Journal of South American Earth Sciences. 11(3):217—231. DeEVRIES, T. J.. L. M. Storr & W. J. ZINSMEISTER. 1984. Neogene forearc basins in southern Chile. Antarctic Journal of the United States 19(2):12-13. DEVRIES, T. J. & G. J. VERMEU. 1997. Herminespina: new genus of Neogene muricid gastropod from Peru and Chile. Journal of Paleontology 71(4):610-615. DiSatvo, L. H. & M. R. CarrikeER. 1994. Planktonic, metamor- phic, and early benthic behavior of the Chilean loco Con- cholepas concholepas (Muricidae, Gastropoda, Mollusca). Journal of Shellfish Research 13(1):57—66. DowsetTT, H., J. BARRON & R. Poore. 1996. Middle Pliocene sea surface temperatures: a global reconstruction. Marine Mi- cropaleontology 27:13—25. Dye, A. H. 1991. Feed preferences of Nucella crassilabrum and juvenile Concholepas concholepas (Gastropoda: Muricidae) from a rocky shore in southern Chile. Journal of Molluscan Studies 57:301—307. HERM, D. 1969. Marines Pliozain und Pleistozan in Nord- und Mittel-Chile unter besonderer Berticksichtigung der En- twicklung der Mollusken-Faunen. Zitteliana 2:159 pp. Hupr, H. 1854. Malacologia y conquiliogia. In: C. Gay (ed.), Historia fisica y politica de Chile. Volume 8 and Atlas (Zool- ogica). Maulde et Renou: Paris. IBARAKI, M. 1992. Neogene planktonic foraminifera of the Ca- mana Formation, Peru: their geologic age and implications. Reports of Andean Studies, Shizuoka University, Special Volume 4:9-19. KENSLEY, B. 1985. The fossil occurrence in southern Africa of the South American intertidal mollusc Concholepas concho- lepas. Annals South African Museum 97(1):1—7. LAMARCK, J. B. 1801. Systeme des Animaux sans Vertebres, ou Tableau General des Classes, des Ordres et des Genres des Ces Animaux. Paris. 432 pp. MARINCOVICH, L., JR. 1973. Intertidal mollusks of Iquique, Chile. Natural History Museum (Los Angeles County) Science Bulletin 16:49 pp. Moreno, C. A., A. Reyes & G. ASENCIO. 1993. Habitat and movements of the recruits of Concholepas concholepas (Mollusca; Muricidae) in the rocky intertidal of southern Chile. Journal of Experimental Marine Biology and Ecology 171:51-61. MuIZoNn, C. DE & T. J. DEVRtES. 1985. Geology and paleontology of the Pisco Formation in the area of Sacaco, Peru. Geolo- gisches Rundschau 74(3):547-563. OLcHAUSKI L., E. 1980. Geologia de los cuadrangulos de Jaqui, Coracora, Chala, y Chaparra. Instituto Geologico Minero y Metalurgico, Boletin 34:71 pp. PEcHOo G., V. & G. Morates S. 1969. Geologia de los cuadran- gulos de Camana y La Yesera. Servicio de Geologia y Mi- neria, Boletin 21:72 pp. Petucu, E. J. 1995. Molluscan diversity in the late Neogene of Florida: evidence for a two-staged mass extinction. Science 270:275—277. Pitsspry, H. A. 1916. The sessile barnacles (Cirripedia) contained in the collections of the U.S. National Museum; including a monograph of the American species. United States National Museum Bulletin 93:366 pp. Soot-RYEN, T. 1959. Pelecypoda. Reports of the Lund University Chile Expedition 1948—49 55(6):3—85. STANLEY, S. M. 1986. Anatomy of a regional mass extinction: Plio-Pleistocene decimation of the western Atlantic bivalve fauna. Palaios 1:17—36. Page 50 TAVERA, J. 1979. Estratigraffa y paleontologia de la Formacion Navidad, Provincia de Colchagua, Chile (Lat. 30°50’S). Mu- seo Nacional de Historia Natural (Santiago, Chile) Boletin 36:176 pp. Watters, W. A., & C. A. FLEMING. 1972. Contributions to the geology and paleontology of Chiloe Island, Southern Chile, parts I and II. Philosophical Transactions of the Royal So- ciety of London (series B) 263:373—407. APPENDIX List of Localities Locality 95DV 809. Yauca, roadcut on west side of Panamerican Highway as it descends to valley floor; 15°39'49"S, 74°31'50”W (Yauca 1:100,000 quadrangle). Locality 9SDV 812. South side of Quebrada Huacllaco, roadcut along Panamerican Highway (15°52’S, 74°11'W, Chala 1:100,000 quadrangle). Locality 95DV 815. Quebrada Chiroteo, 0.7 km north of juncture with Quebrada Mal Paso; roadcut on east side of the Panamerican Highway, at an elevation of about 520 The Veliger, Vol. 43, No. 1 m.s.1., 16°34'41"S, 72°37'33”W (Camana 1:100,000 quad- rangle). Locality 9SDV 816. Quebrada Chiroteo, 0.5 km north of juncture with Quebrada Mal Paso; roadcut on east side of the Panamerican Highway, at an elevation of about 500 m.s.1., 16°34'48"S, 72°37'32”W (Camana 1:100,000 quad- rangle). Locality 96DV 911. Acari Depression, 6 km north- northeast of Chavifia, southeastern base of ridge extend- ing eastward into depression, 15°34'37"S, 74°36'20”W (Yauca 1:100,000 quadrangle). Locality 96DV 923. About 0.5 km southeast Monte Redondo, east side Rio Acari, ridge inside shallow, small depression, 15°36'29"S, 74°37'53”W (Yauca 1:100,000 quadrangle). Locality 97DV 935. Northwest side of Cerro Antivo, south of road between Pozo Santo and Laguna Grande, at pass southwest of Cerro El Diablo. Inlier of bioclastic sandstone and gravel. 14°03’49"S, 76°08'58”"W (Punta Grande 1:100,000 quadrangle). The Veliger 43(1):51—57 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Pisidium taraguyense and Pisidium pipoense, New Species from Northeastern Argentina (Bivalvia: Sphaeriidae) CRISTIAN FE ITUARTE Departamento Zoologia Invertebrados, Museo de La Plata, 1900 - La Plata, Buenos Aires, Argentina (e-mail: cituarte@museo.femym.unlp.edu.ar) Abstract. Pisidium taraguyense sp. nov., from Corrientes province, and Pisidium pipoense sp. nov., from Misiones province, northeastern Argentina, are described. Pisidium taraguyense sp. nov. is characterized by a definite ovate shell outline, markedly inequilateral shell, quite inflated, with beaks full, ligament tending to be external, hardly visible from the exterior, and never protruding. Pisidium pipoense sp. nov. is characterized by a subquadrangular shell outline, with inflated and centrally located beaks, ligament tending to be external, visible from the exterior, but not protruding. In both species, the roof of the ligament-pit is a very thin wall formed by a series of small calcic plates that partially hide the ligament, which is exposed (only at its anterior half) through a narrow line. INTRODUCTION Little is known about the species of Pisidium Pfeiffer, 1821, distributed in Misiones and Corrientes provinces (Figure 1), a geographical area characterized from a fau- nistic viewpoint by the high degree of local and regional endemism evidenced by several freshwater mollusks e.g., the monospecific genus Acrorbis Odhner, 1937 (Basom- matophora: Planorbidae) (Paraense, 1986); one species of Chilina Gray, 1828 (Basommatophora: Chilinidae) (Hyl- ton Scott, 1958); one species of Pomacea Perry, 1810 (Mesogastropoda: Ampullariidae) (Ihering, 1919), and two species of Eupera Bourguignat, 1854 (Bivalvia: Sphaeriidae) (Ituarte, 1989; Ituarte & Mansur, 1993). Knowledge of the taxonomy of the genus Pisidium in southern South America is also rather scarce, and it main- ly comes from works of d’Orbigny (1835, 1846); Pilsbry (1897, 1911); Meier-Brook (1967); Kuiper & Hinz (1984); Ituarte & Gordillo (1991); and Ituarte (1995, 1996). In the present paper, the presence of the genus Pisidium in Misiones and Corrientes provinces is reported for the first time, and two new species, Pisidium pipoense and Pisidium taraguyense, are also described. MATERIALS anD METHODS Samples upon which the present study was based are part of the malacological collection of the Department of In- vertebrate Zoology, Museo de La Plata (MLP). Addition- al samples were taken by the author during the years 1987, 1991, and 1997, now lodged in the same institution. Paratypes of Pisidium forense Meier-Brook, 1967 (from Senckenberg Museum, Frankfurt [SMF]) were studied for comparative purposes. Length measurements (shell length [SL], shell height [SH], shell width [SW], and pre-si- phonal suture length [PSS]), shape indices and morpho- metric ratios (height index [I SH/SL], convexity index [Ci = SW/SH], and hinge length: shell length ratio [HiL/ SL]), were calculated according to the criteria followed by Ituarte (1996). SYSTEMATICS Pisidium taraguyense Ituarte, sp. nov. (Figures 2—10) Diagnosis: Medium-sized species characterized by ovate shell, rather inequilateral, with anterior end produced, quite inflated with full beaks. Ligament tending to be external but hardly visible from the exterior, never protruding. Description: Species medium-sized (maximum SL = 4.1 mm), quite inflated (average Ci = 84.09 + 3.6), rather inequilateral. Beaks full, wide, well visible above dorsal margin, backwardly displaced (located at about 62% of shell length). Shell outline sharply ovate, sometimes tend- ing to be elliptical; however, due to prominent beaks above dorsal margin, height index (shell height/shell length) does not reflect actual appearance of shell outline (average I SH/SL = 84.1 + 2.53). Dorsal margin mark- edly shorter than ventral one, very slightly concave at midst, anterior half straight, posterior one evenly curved. Anterior margin projecting in sharp curve, posterior mar- gin rounded, somewhat truncated. Well-marked angle be- tween dorsal and anterior margin. Ventral margin wide, evenly curved, somewhat straight in cases. Shell surface dull, light yellowish brown color, rather regularly and finely striated (more than 40 striae per mm). Hinge plate quite narrow, left inner (C,) and right (C;) cardinal teeth hanging from inner hinge margin. Hinge line markedly curved, rather long (HiL/SL = 51.6 + 1.6). Manantia ° J BUENOS ~= Figure | Location map. In left valve, two delicate cardinal teeth, the internal (C,) very slightly sinuous, upward directed at posterior end; external cardinal tooth (C,) slender, nearly straight, slight- ly oblique, located just below central point of beak, over- lapping C, at its posterior half or slightly more. Anterior lateral tooth (Ay) close to cardinals, strong, triangular cusp displaced forward. Posterior lateral tooth (P,,) short, robust, with distally displaced cusp. Right valve: a deli- cate cardinal tooth (C;), extremely slender at anterior half; straight, enlarged at posterior end, forming triangular, weakly sulcated small head. Anterior lateral teeth well developed, inner (A,) with distal cusp, outer (Ay) a mi- nuscule short triangular cusp; outer posterior lateral tooth (Py) short, triangular, low, with nearly central cusp, inner one (P,) with distally displaced triangular cusp. Diver- gence angle between laterals about 130°. Ligament slightly tending to be external, not protrud- ing, hardly discernible from exterior. Ligament-pit long, low, and slender (representing about 23% of shell length); The Veliger, Vol. 43, No. 1 Figures 2 and 3 Pisidium taraguyense Ituarte, sp. nov. (MLP 5318). Figure 2. Right external view. Figure 3. Posterior view of a shell. Scale bar (same scale bar for both figures) = 2 mm. partially closed dorsally by very thin calcareous wall, ac- tually formed by several (four to six) small plates (easily detachable in dried specimens); then, ligament visible from exterior only from a narrow line. Anatomy: Only one siphonal aperture, the anal. Pre-si- phonal suture length about 15% of shell length, nearly equal to diameter of anal aperture. Only inner demi- branchs present. Brood pouches arising from upper and posterior part of demibranchs. Remarks: Pisidium taraguyense differs from the remain- ing Argentinian Pisidium species in being decidedly ovate, or elliptical, in shell outline, strikingly inequilater- al, and quite globose. It shares with Pisidium sterkianum Pilsbry, 1897, and Pisidium vile Pilsbry, 1897 (both spe- cies distributed along the Parana, Rio de la Plata and Uru- guay basins) the presence of only one siphonal aperture and only one, the inner, demibranch as common features. The external ligament is also a common characteristic; however, in Pisidium taraguyense it is not as extroverted as in P. sterkianum. The same type of small extroverted external ligament has also been observed in Pisidium pi- C. F. Ituarte, 2000 Page 53 poense (present study), and in the studied paratypes of Pisidium forense Meier-Brook, 1967 (SMF 186492/c and 186492/h) from Juiz de Fora, Minas Gerais, Brazil. Meier-Brook (1967) reported that in some paratypes of P. forense the ligament seems to be completely enclosed. Type locality: La Cruz (29°10'S, 56°38'W), San Martin department, Corrientes Province, Argentina. Distribution: Known only from Corrientes Province, be- sides the type locality, at Mocoreta (unnamed brook) (30°28'S, 58°04’W) (MLP 5347), Monte Caseros depart- ment, and Manantiales (27°56’S, 58°06’W) (MLP 5317), Mburucuya department (Figure 1). Type specimens: Holotype: in the collection of the De- partment of Invertebrate Zoology, Museo de La Plata (MLP 5318); paratypes: MLP 5318; Division of Inverte- brates, Museo Argentino de Ciencias Naturales (2 para- types MACN 34120); Muséum National d’Histoire Na- turélle (Paris) (3 paratypes). Etymology: The specific name derives from Taraguy (or Taragtii), the aboriginal name given by the Guarani Indian people to Corrientes city. The meaning of the name was later extended by usage to designate the whole territory that presently corresponds to Corrientes Province, where the type specimens were collected; it was part of the vast ancient territory occupied by the Guarani tribes. Pisidium pipoense Ituarte, sp. nov. (Figures 11—19) Diagnosis: Shell rather solid, small or medium-sized, in- flated, beaks nearly central. The subquadrangular shell outline is the distinctive character. Ligament slightly tend- ing to be external, visible from the exterior but not pro- truding. Description: Shell small to medium (maximum SL = 3.9 mm), rather solid, inflated (average CI = 74.19 + 5.44), nearly equilateral. Shell outline sub-quadrangular, high (average I SH/SL = 89.2 + 1.68), dorsal margin slightly shorter than ventral margin. Anterior half of dorsal mar- gin straight, posterior half, slightly convex. Ventral mar- gin evenly curved. Anterior end moderately produced, posterior end truncated. Beaks full, nearly central (located at about 58% of shell length), well projecting above dor- sal margin, slightly opistogyrous. Nepionic shell usually Figures 4—7 Pisidium taraguyense Ituarte, sp. nov. (MLP 5318). Figure 4. Hinge of the left valve. Figure 5. Detail of left cardinal teeth, ligament, and posterior lateral tooth. Figure 6. Hinge of the right valve. Figure 7. Detail of right cardinal tooth, ligament, and pos- terior lateral teeth. Scale bars = 1 mm. Page 54 Figures 8—10 Pisidium taraguyense Ituarte, sp. nov. (MLP 5318). Figure 8. Lateral view of the ligament position and dorsal wall (arrow) of the ligament-pit. Figure 9. Postero-dorsal view of the escutcheon, showing the calcic plates forming the roof of the ligament-pit, and the narrow part of the ligament exposed (arrow). Figure 10. Dorsal view of the escutcheon, showing the space left by the lack of two or three plates detached during the processing of the specimen for SEM (arrow). Scale bars: Figure 8-9 = 1 mm; Figure 10 = 200 pm. The Veliger, Vol. 43, No. 1 Figures 11 and 12 Pisidium pipoense Ituarte, sp. nov. (MLP 5336). Figure 11. Right external view. Figure 12. Posterior view of a shell. Scale bar (same scale bar for both figures) = 1 mm. marked, forming a cap. Surface finely and regularly stri- ated (about 30—40 striae per mm). Hinge plate somewhat solid. Hinge line long (HiL/SL = 52 + 2.1). In right valve, a single, somewhat weak cardinal tooth (C;), slender, low, faintly curved, enlarged at posterior end, forming a slightly sulcated cup. Anterior right lateral teeth well developed, the outer (Aj,) small, cusp slightly displaced forward, the inner (A,) robust, cusp central. Outer posterior lateral tooth (P,;,) reduced in size, cusp distal, forwardly displaced with respect to P,. P, strong, cusp triangular, central. In left valve, two well- developed cardinal teeth. Inner cardinal tooth (C,) short, somewhat bent upward at distal end, located before cen- tral point of beak. Outer left cardinal (C,) long, slender, slightly arcuate, parallel with respect to antero-posterior axis, overlapping C, at nearly its entire length. Anterior (A,) and posterior (P,,) lateral teeth strong, both arising from robust bases, cusps high, triangular, distally dis- placed in A, and nearly central in P,. Divergence angle between laterals varies from 115° to 120°. Ligament visible from exterior only as narrow line, never protruding. Ligament-pit slender-and long, surpass- C. F. Ituarte, 2000 Page 55 ing (sometimes largely) the interception line of beak with dorsal margin (ligament length about 21% of shell length). Dorsally, ligament-pit delimited by very thin cal- careous wall. In dried specimens it breaks, usually into a series of four to six small delicate, parallelogram-shaped calcic plates. This thin wall almost closes completely lig- ament-pit, determining when valves are joined; only a very narrow space remains, from which ligament is ex- posed to exterior. Anatomy: Only anal aperture present. Pre-siphonal su- ture long, representing about 20% of shell length, and 1.6 of diameter of siphonal aperture (average 1.55 + 0.31). Only inner demibranch present; brood pouch develops from upper part of inner wall of descendent lamella. Five well-marked muscle scars, corresponding to inner radial mantle muscles, located far from pallial line. Remarks: Pisidium pipoense may be easily distinguished from other South American Pisidium species by its gen- eral external shape and subquadrangular shell outline. P. pipoense is similar to Pisidium forense Meier-Brook, 1967, but is distinguished by its smaller maximum size, slightly lower and flatter shell, sharp quadrangular shell shape, and stronger hinge plate. The more central position of the beaks and the shape and relative length, slightly longer, of the ligament are also distinctive features. The shell surface striation is more marked, fine, and regularly spaced in P. forense than in P. pipoense. Beaks are fuller in P. forense, and the presiphonal suture is longer than in P. pipoense. In P. pipoense the position of the liga- ment, tending slightly to be external (never protruding), the presence of only one siphonal aperture and one (the external) demibranch, and the type of development of marsupial sacs (arising upwardly and posteriorly in inner demibranchs) resemble species belonging to the “‘eastern Brazilian”’ and ‘‘Parano-Platense”’ drainages (Figure 1) (i.e., Pisidium sterkianum and Pisidium vile from Uru- guay and Argentina, and Pisidium forense from Brazil). However, the presence of a thin wall partially closing the roof of the ligament-pit, as was observed in P. forense and P. taraguyense, and the somewhat solid hinge plate are substantial differences with respect to the only Ar- gentine species known to have external ligaments: P. ster- kianum. Type locality: A small creek that flows into the Santo Pip6 Brook (27°07'S, 55°24’W), on the intersection with Figures 13-16 Pisidium pipoense Ituarte, sp. nov. (MLP 5336). Figure 13. Hinge of the right valve. Figure 14. Detail of right cardinal tooth, ligament, and posterior lateral teeth. Figure 15. Hinge of the left valve. Figure 16. Detail of left cardinal teeth, ligament, and pos- terior lateral tooth. Scale bars = 1 mm. Page 56 Figures 17-19 Pisidium pipoense Ituarte, sp. nov. (MLP 5336). Figure 17. Lat- eral view of the ligament position and dorsal wall (arrow) of the ligament-pit. Figure 18. Latero-dorsal view of the escutcheon showing the calcic plates forming the roof of the lgament-pit (arrow). Figure 19. Dorsal view of the escutcheon, showing the narrow exposed area of the ligament (arrow). Scale bars = 100 p.m. the national route No. 12, Santo Pip6, department of San Ignacio, Misiones, Argentina. Distribution: Known only from the type locality. Type specimens: Holotype: in the collection of the De- The Veliger, Vol. 43, No. 1 partment of Invertebrate Zoology, Museo de La Plata (MLP 5336); paratypes: MLP 5298 and 5336; Division of Invertebrates, Museo Argentino de Ciencias Naturales (2 paratypes MACN34119); Muséum National d’ Histoire Naturélle (Paris) (3 paratypes). Etymology: The specific name refers to the type locality, Santo Pip6o. GENERAL REMARKS Our limited knowledge of the specific diversity and mor- phological variability of the sphaeriid fauna from South America precludes us from proposing natural supraspe- cific groups for Pisidium species distributed in the Neo- tropical Region. However, new data available from the descriptions of Pisidium taraguyense and Pisidium pi- poense allow us to envisage at least a preliminary bio- geographical sketch for a restricted geographical area in eastern South America, within which seems to arise a somewhat clearly defined group of species distributed along the drainages of southeastern Brazil, western Uru- guay, and northeastern Argentina, with a provisional southern limit in the Rio de la Plata River (Figure 1!) (i.e., the southern limit of the known geographical distribution of P. vile and P. sterkianum (Ituarte, 1996). This group of species includes P. sterkianum, P. vile, P. forense, P. taraguyense, and P. pipoense. The morphological basis for this group, still incomplete, includes the possession of a fragile shell, a rather weak hinge plate, only one de- mibranch and siphonal opening, as well as any sort of external ligament, ranging from a true external ligament, as in P. sterkianum, to an external ligament partially hid- den by the development of the dorsal wall of the liga- ment-pit and never protruded as in P. forense, P. tara- guyense, and P. pipoense. The species included in this group seem to differ consistently from those species dis- tributed in southern Chile and Patagonia (Argentine ter- ritories south to the Colorado River), which have more solid shells and hinge plates, an internal ligament and, in some, two demibranchs and two siphonal apertures (Pils- bry, 1911; Ituarte, 1996). Acknowledgments. Dr. Ronald Janssen (Forschungsinstitut und Naturmuseum Senckenberg) kindly contributed to this study by providing paratypes of Pisidium forense. The invaluable support of Mr. Luis Biestro, Dr. Gustavo Darrigran, Dr. Maria Pujals, and Santiago Ituarte during field trips; and the fine work done by Lic. Rafael Urréjola (scanning electron microscopy unit MLP) is also acknowledged. The author is a researcher of the Conselo Na- cional de Investigaciones Cientificas (CONICET), Argentina. LITERATURE CITED D’ORBIGNY, A. D. 1835. Synopsis terrestrium et fluviatilium mol- luscorum, in suo per American meridionale itinere, ab A, d’Orbigny, collectorum. Magasin de Zoologie 6(61—62):1— 44. D’ORBIGNY, A. D. 1835-1846. Voyage dans l’Amerique Méri- C. F. Ituarte, 2000 dionale exécuté pendant les années 1826-1833. Vol. 5, part. 3, Mollusques. ed. P. Bertrand: Paris. HyLTon Scott, M. I. 1958. Nueva especie de Chilina del norte argentino (Moll. Pulm. Basommatophora). Neotropica 4(13): 26-27. THERING, H. 1919. Las especies de Ampullaria de la Argentina. Primera Reunion Nacional de la Sociedad Argentina de Ciencias Naturales, seccion 4, Zoologia:329—350, 2 pls. ITUARTE, C. F 1989. Los géneros Byssanodonta d’Orbigny, 1846 y Eupera Bourguignat, 1854 (Bivalvia: Sphaeriidae) en el area Parano-Platense. Descripcion de Eupera iguazuensis n. sp. del rio Iguazu, Misiones, Argentina. Neotropica 35(93): 53-63. ITUARTE, C. E 1995. Nuevos registros de Pisidium Pfeiffer, 1821 y Sphaerium Scopolhi, 1777 (Bivalvia: Sphaeriidae) en Chile, Bolivia y Noroeste argentino. Neotropica 41(105—106):31— 41. ITUARTE, C. F 1996. Argentine species of Pisidium Pfeiffer, 1821, and Musculium Link, 1807 (Bivalvia: Sphaeriidae). The Ve- liger 39(3):189-—203. Page 57 ITuaRTE C. F & S. GORDILLO. 1991. Nuevas citas de pelecipodos dulciacuicolas de Isla Gable, Tierra del Fuego, Argentina. Neotropica 37(97):29—30. ITUARTE, C. EF & M. C. Mansur. 1993. Eupera elliptica n. sp. (Bivalvia: Sphaeriidae), una nueva especie en el rio Iguazu, Misiones, Argentina. Neotropica 39(1):11—16. KurpPer, J. G. J. & W. Hinz. [1983] 1984. Zur fauna der klein- muscheln in den Anden (Bivalvia: Sphaeriidae). Archiv fiir Molluskenkunde 114(4—6):137—156. MEIER-BROOK, C. 1967. Pisidium forense, a new species from Brazil (Mollusca; Eulamellibranchiata; Sphaeriidae). Archiv fiir Hydrobiologie 64(1):63-68. PARAENSE, W. L. 1986. The radula of Acrorbis petricola (Pul- monata: Planorbidae). The Nautilus 100(3):109—112. Pitssry, H. A. 1897. New species of mollusks from Uruguay. Proceedings of the Academy of Natural Sciences of Phila- delphia, May 1897:290—298, 2 pls. Pitssry, H. A. 1911. Non-marine Mollusca of Patagonia. Reports of the Princeton University Expedition to Patagonia (1896— 1899), 3(5):513-633. The Veliger 43(1):58—63 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Development and Metamorphosis of the Planktotrophic Larvae of the Moon Snail, Polinices lewisii (Gould, 1847) (Caenogastropoda: Naticoidea) ROBERTA V. K. PEDERSEN AnD LOUISE R. PAGE* Department of Biology, University of Victoria, RO. Box 3020 STN CSC, Victoria, British Columbia, Canada V8W 3N5 Abstract. Polinices lewisii from the east coast of Vancouver Island, British Columbia, has been reported previously to have short-lived, non-feeding larvae and juveniles that feed on diatoms and Ulva sp. before switching to bivalve prey. However, we found that egg masses of this species from three sites along the southeast and west coast of Vancouver Island, British Columbia, released planktotrophic larvae. Larvae fed Isochrysis galbana could be induced to metamor- phose at 5 weeks after hatching when cultured at 20—22°C, but repeated attempts to rear larvae to metamorphic com- petence at 12°C were unsuccessful. Major events of larval development include: bifurcation of the velar lobes, elongation of cephalic tentacles, great enlargement and elaboration of the foot, growth of the larval shell to 2% whorls, accumulation of black pigment within the epidermis and wall of the gut, and differentiation of the osphradium and gill lamellae. Metamorphosis was induced by sediment from an embayment inhabited by adults, but the active factor was destroyed by autoclaving. Juvenile bivalves had minimal inductive potency. Beginning at 3 to 5 days after metamorphic velum loss, P. /ewisii drilled and ingested both juvenile bivalves and ostracods; we did not find evidence of feeding on Ulva. INTRODUCTION Members of the gastropod superfamily Naticoidea are predators of infaunal bivalves and gastropods (see review by Kabat, 1990), which they drill by radular rasping and corrosive secretions from an accessory boring organ at the tip of the acrembolic proboscis (see review by Car- ricker, 1981). Polinices lewisii (Gould, 1847) is the larg- est extant member of this cosmopolitan caenogastropod superfamily (Marincovich, 1977). The species inhabits fine sediment embayments from the low intertidal to shal- low subtidal zone along the west coast of North America. The feeding ecology and digestive physiology of P. lew- isii have been studied extensively (Bernard, 1967; Reid & Freisen, 1980; Reid & Gustafson, 1989; Peitso et al., 1994). Naticids are also notable for the large size of their foot, which is used for burrowing through surface sedi- ments and for capture and subjugation of prey (Ziegel- meier, 1954, 1958; Fretter & Graham, 1962:572-574; Bernard, 1967; Hughes, 1985). The foot of P. lewisii can be inflated to four times the shell volume by uptake of seawater into channels within the foot (Bernard, 1968). Naticid females deposit their internally fertilized eggs within a benthic egg mass, which is usually a collar- shaped conglomerate of egg capsules and sand held to- gether by a rubbery adhesive (an exception is described by Booth, 1995). Depending on the species, progeny hatch as: (1) swimming, planktotrophic larvae; (2) swim- * Corresponding author: Phone: (250) 472-4679; Fax: (250) 721-7120; e-mail: lpage@uvvm.uvic.ca ming, lecithotrophic larvae (short-lived relative to the planktotrophs); or (3) crawl-away juveniles (Thorson, 1935; Giglioli, 1955; Amio, 1955; Bernard, 1967). Oc- casionally, two different life history patterns have been reported for a single naticid species. Thorson (1935) re- ported direct development for Natica (Polinices) catena (Da Costa, 1778), a species that ingests nurse eggs during embryological development, but Lebour (1936) described swimming larvae for this species. Giglioli (1955) re- viewed evidence that Polinices triseriata Say, 1826, may hatch as either crawl-away juveniles or swimming veli- gers. Finally, and of direct relevance to our study, Ber- nard (1967) reported short-term, lecithotrophic larvae for P. lewisii, but Giglioli (1955) predicted planktotrophic larvae on the basis of embryo and egg capsule dimen- sions measured from preserved egg masses. Our observations on the life history of P. lewisii con- firm Giglioli’s (1955) prediction of planktotrophic larvae. This result is based on egg masses collected from one location on the west side and two locations on the south- east side of Vancouver Island, British Columbia, Canada. Collections were made during 4 successive years from one of the latter locations. In this report, we describe general aspects of development from larval hatching through metamorphosis into young juveniles. MATERIALS AND METHODS Collection and Maintenance of Egg Masses We examined larvae of Polinices lewisii that hatched from egg masses laid in the following localities and years: R. V. K. Pedersen & L. R. Page, 2000 (1) Barkley Sound (west coast of Vancouver Island), sum- mer 1995; (2) Bamberton Harbour, Saanich Inlet (east coast of southern Vancouver Island), summer 1997; and (3) Patricia Bay, Saanich Inlet, summer 1994 and spring to late summer 1995, 1996, 1997. Egg masses were col- lected during low tides from muddy sand beaches where adults were also found. Egg masses were maintained in aquaria provided with flowing seawater from the recir- culating system at the University of Victoria and were aerated continuously from a compressed air outlet until larvae began to emerge. Larval Culture Larvae were reared in glass flasks containing 500 mL of seawater that was coarse filtered under vacuum with a Millipore prefilter (HAP 20 047 00). The unicellular alga, Isochrysis galbana, was added to the larval cultures at a concentration of 5 X 10* cells/mL for the first 2 weeks and 10° cells/mL thereafter. Isochrysis galbana was cul- tured in filter-sterilized seawater enriched with nutrient medium purchased from Fritz Chemical Company (Dal- las, Texas). Algal cells were washed before they were added to the larval cultures by centrifuging aliquots of algal culture at approximately 1000 RPM for 7 min, dis- carding the supernatant, and resuspending the algal pellet in coarse-filtered seawater. Density of algal cells was de- termined with a hemacytometer. Young larvae were cul- tured at an initial density not exceeding one larva per 2 mL seawater, but this was gradually reduced by subdi- viding cultures and removing larvae until a density not exceeding one larva per 10 mL was reached by 3 weeks post-hatching. Larvae were transferred to fresh culture medium every 2 or 3 days by a combination of gentle sieving and hand pipetting. Seawater for culturing larvae was collected once weekly from a rocky coastal prom- ontory in an area subject to strong tidal mixing. It was stored at 12°C in Nalgene carboys and coarse filtered im- mediately before use. Larvae were cultured at either 12°C or 20—22°C. Light and Scanning Electron Microscopy To prevent larvae from retracting into the shell during light microscopical examination and photography, they were anaesthetized in artificial seawater having excess Meg?* and reduced Ca** (Audesirk & Audesirk, 1980) for periods ranging from 2 hours for hatching larvae to 24 hours for older larvae and juveniles. The cleaned larval shell shown in Figure 1f was obtained by placing anaes- thetized larvae in distilled water for 30 minutes, followed by a 6-hour immersion in a 3:10 mixture of household bleach in distilled water. To remove residual, lipid-rich tissues, the shells were taken gradually into 100% meth- anol, then into a 1:1 mixture of methanol and chloroform for 0.5—1 hr, then back through the methanol series to distilled water. Page 59 Larval shells were prepared for scanning electron mi- croscopy according to the method of Hadfield & Strath- mann (1990). Cleaned veliger shells and ostracod cara- paces drilled by juvenile P. /Jewisii were stored in absolute acetone. They were subsequently air dried, mounted on stubs using nail polish as an adhesive, sputter coated with gold, and photographed with a JEOL SM35 scanning electron microscope. RESULTS Larval Development All egg masses of Polinices lewisii that we collected released swimming larvae (Figure la, b). When recently hatched larvae were examined microscopically at 1 hour after being placed in seawater containing /sochrysis gal- bana at 5 X 10* cells/mL, algal cells were seen within the larval stomach. Four attempts to rear larvae at 12°C failed to produce metamorphically competent veligers af- ter 3% months, when the cultures were terminated. How- ever, most larvae cultured at 20 to 22°C were competent to metamorphose at 5 weeks after hatching, some as early as 4 weeks. Larvae that could use their foot for crawling were judged to be competent to metamorphose. Compe- tent larvae tended not to crawl on a clean glass surface, but when cultures were poured through a sieve during the procedure for transferring larvae to fresh culture medium, competent larvae often crawled on the Nitex cloth (64 jzm mesh opening) used to construct the sieves. The fol- lowing description of the pattern and timing of develop- mental events for P. lewisii is based on larvae that were reared at 20—22°C. Hatching larvae of P. lewisii had a bilobed velum, a small foot and shell, and a functional digestive tract (Fig- ure la, b). A pre-oral and post-oral ciliary band, with an intervening tract of food groove cilia, extended around the periphery of each velar lobe. The shell at hatching stage (embryonic shell or protoconch I) consisted of slightly more than one whorl, had a punctated surface sculpture (Figure Ic), and was almost bilaterally sym- metrical. When measured in lateral view from the outer lip of the aperture to the opposite side of the shell, pro- toconchs of hatching larvae had a mean value of 235.4 pm (SD 5.5 wm; n = 5). Rudiments of the cephalic ten- tacles were recognizable as a pair of stubby papillae at the apex of the larval body, and pigmented eyespots were visible through the transparent cephalic epidermis (Figure la). The larval heart was a thin-walled vesicle located beneath the floor of the mantle cavity that showed rhyth- mic contractions in hatching larvae, although contractions stopped when the foot and velum retracted into the shell. Occasional bouts of repetitive contractions in the area of the definitive heart were also evident in hatching larvae. These occurred slightly posterior to the larval heart. The strength and regularity of definitive heart contractions in- creased during the days following hatching, but the pe- Page 60 The Veliger, Vol. 43, No. 1 Figure | Larval and metamorphic development of Polinices lewisii. a. Newly hatched larva in antero-ventral view showing the two disc-shaped velar lobes (VL), shell (SH), eye (E), mouth (M), and statocysts (S); scale bar, 75 wm. b. Newly hatched larva in right lateral view showing the small foot (F) and operculum (QO); the stomach (ST) and left digestive R. V. K. Pedersen & L. R. Page, 2000 riodicity was not synchronous with that of the larval heart. Both the soft tissues and shell showed substantial growth between hatching and metamorphic competence (compare Figure 1b and g). Each of the enlarging velar lobes acquired two red pigment spots by 6 days post- hatching, and by 9 days each velar lobe began to bifurcate (Figure 1d, e). Eventually, the velum consisted of four long arms with a patch of red pigment embedded in both the upper and lower velar epidermis at the apex of each arm (Figure 1g). Formation of a four-armed velum and subsequent lengthening of each velar arm greatly in- creased the length of the velar ciliary bands relative to the initial bilobed condition. Shell secreted during the larval stage (larval shell or protoconch II) had incremental growth lines but no sur- face punctae (Figure lc). The growing larval shell spi- raled in the orthostrophic direction (torsional asymmetry was dextral), which generated a low spire on the right and an umbilicus on the left (Figure 1f). At metamorphic competence, the shell had 2% whorls and a mean diameter of 941 pm (SD 41 pm; n = 7) when measured from the outer lip of the aperture to the opposite side of the shell. The foot of hatching larvae was a small, triangular ex- tension of post-oral body wall that bore the operculum. At this initial larval stage, the foot had a low dorso-ven- tral profile along its entire length and lacked pigmentation (Figure 1b). The metapodial region of the foot began to spread laterally soon after hatching, but the most dramatic change in pedal form during the first third of larval life was the expansion of the propodium (Figure le). By 2 weeks post-hatching, the propodium was almost worm- like in shape and flexibility. After 2 weeks, the foot ac- quired the beginnings of the propodial and metapodial folds. Following metamorphosis, these folds enlarge suf- ficiently to almost completely cover the external surface of the shell; they function to prevent sediment from en- tering the mantle cavity during post-metamorphic bur- rowing behavior. As the foot enlarged, there was a gradual accumulation of black pigment within the originally transparent pedal epidermis. Black pigment also appeared within the wall of the gut, within the epidermis of the head (excluding the velum), and within the floor of the mantle cavity. The mantle fold was sparsely pigmented, except for a strip of Page 61 black that ran parallel to the developing osphradium. The black pigment strip became obvious approximately 3 weeks after hatching. Other notable developmental events that we saw in whole mounts of live larvae included the elongation of cephalic tentacles, which occurred throughout the larval phase, and the appearance of gill lamellae along the roof of the mantle cavity at approximately 3 weeks post-hatch- ing. After approximately 5 weeks of laboratory culture, ve- ligers of P. lewisii acquired the ability to crawl over sur- faces or burrow into fine sediments using the darkly pig- mented foot. When crawling began, the velar arms were retracted into the shell, and the remarkable foot of these late-stage veligers became greatly inflated so as to form a broad ventral platform beneath the shell. The inflation, together with the contours of the propodium, gave the leading face of the crawling foot a shovel-like shape (Fig- ure lh). As this highly maneuverable appendage was thrust into sediment during burrowing, cilia along the frontal slope of the foot generated a conveyor-belt-like transport of sediment particles from immediately in front of the foot, along the anterior face of the propodium, to the dorsal surface of the shell. The foot deflated rapidly and reverted to a shrunken condition when veligers stopped crawling or burrowing and either retracted fully into the shell or resumed swimming (Figure 1g). Metamorphosis Although we reared greater than 200 larvae of P. lew- isii to metamorphic competence, some for a month past the onset of crawling ability, we identified only two cases of spontaneous metamorphosis in glass culture flasks con- taining coarse-filtered seawater. However, approximately 75% of larvae that were able to crawl underwent meta- morphosis when placed in bowls of seawater containing approximately 1 cc of surface sediment from an area in- habited by adults (Patricia Bay site). Sediment that was autoclaved, then rinsed in filter-sterilized seawater, com- pletely lost its inductive capacity. Pieces of Ulva were also ineffective for inducing metamorphosis of this gas- tropod. Small bivalves showed a modest capacity to in- duce metamorphosis of P. lewisii; three of 15 larvae metamorphosed in the presence of small bivalves of gland (LDG) are visible through the transparent shell; scale bar, 75 wm. c. Scanning electron micrograph of the shell from a young larva; arrowhead indicates the boundary between protoconch I and II; scale bar, 50 wm. d. Larva at 11 days post-hatching showing the propodium (PR) and initial bifurcation of velar lobes (VL); scale bar, 100 wm. e. Larva at 11 days post-hatching in left lateral view showing the enlarging metapodium (MP) and propodium (PP) of the foot; scale bar, 100 ym. f. Cleaned shell of a metamorphically competent larva showing 2” shell whorls and orthostrophic coiling direction; scale bar, 250 wm. g. Metamorphically competent larva in right lateral view showing four long velar arms (VL), elongate cephalic tentacles (T), and large foot (F); scale bar, 250 ym. h. Young juvenile in antero-lateral view showing the shovel-like shape of the anterior face of the expanded foot; scale bar, 250 ym. i. Ostracod carapace drilled by a young juvenile of P. lewisii; scale bar, 100 wm. Page 62 mixed species composition that were isolated from Patri- cia Bay sediment. Post-metamorphic individuals could be easily distin- guished from crawling larvae because, during crawling behavior, the cephalic tentacles projected beyond the rim of the shell aperture only after the velar arms had been lost. The mechanism that destroys the velum was not ob- served directly because the velar arms were fully retract- ed into the mantle cavity during metamorphosis. How- ever, we never observed discarded chunks of velar tissue or dissociated velar ciliated cells after competent larvae had been placed in sediment containing the inductive cue. Between 3 to 5 days after loss of the velum, meta- morphosed snails began feeding by drilling holes in shells of small, juvenile bivalves. Juvenile snails appeared to have an adhesive structure at the extreme posterior end of the foot because we occasionally saw juveniles crawl- ing through sediment with a small bivalve attached to this area of the foot. Ostracods were abundant in the sediment samples that we used to induce metamorphosis of P. lew- isii larvae, and these were also drilled and eaten by young, juvenile moon snails (Figure 11). DISCUSSION The literature gives conflicting reports for the develop- mental pattern of Polinices lewisii. Giglioli (1955) pre- dicted planktotrophic larvae for this species, based on embryo and capsule dimensions for egg masses collected from southeastern Vancouver Island at Ladysmith Har- bour. However, Bernard (1967) reported that short-term, non-feeding larvae hatched from egg masses of P. lewisii collected from a slightly more northern site along eastern Vancouver Island at Departure Bay. Bernard’s (1967) ob- servations would seem more credible, since he actually observed hatching larvae of this species. He provided a sketch of an early, encapsulated veliger of P. lewisii, but did not illustrate hatching veligers. Nevertheless, Bernard (1967) stated that hatching P. lewisii larvae had a small velum but a large foot. We found that P. lewisii egg masses collected from three locations around Vancouver Island (collections made during 4 successive years for the Patricia Bay site) released planktotrophic larvae. There are three possible explanations for this discrepancy in reports (1) Bernard (1967) was mistaken; (2) P. lewisii is capable of poecil- ogony (although a number of previous reports of poecil- ogony among gastropods have been disputed by Hoag- land & Robertson [1988] and Bouchet [1989]); or (3) P. lewisii encompasses cryptic sister species that are distin- guishable by a difference in developmental pattern only. The last possibility is not without precedent. Oliverio (1996) reviewed a number of such cases for caenogastro- pods from Europe. We hesitate to overthrow Bernard’s (1967) interpretation because he describes a large foot for hatching larvae of P. lewisii, with the propodium devel- The Veliger, Vol. 43, No. 1 oping within 24 hours. “‘Large’’ would not be an appro- priate adjective for the foot of hatching larvae of P. lew- isii that we observed (Figure 1b). Unfortunately, the site where Bernard (1967) collected P. /ewisii egg masses has been destroyed by dredging for construction of a marine docking facility (Dr. Mary Arai, Pacific Biological Sta- tion, Nanaimo, British Columbia, personal communica- tion). Nevertheless, it would be appropriate to intensively sample populations of P. /ewisii along the west coast of British Columbia to test the hypothesis of cryptic sister species. Our repeated failure to culture larvae of P. lewisii to metamorphic competence at 12°C was unexpected be- cause this is within the normal summer temperature range for marine waters around southern Vancouver Island. At 3% months, larvae cultured at 12°C appeared to be de- velopmentally arrested at a stage corresponding to 60 to 70 percent completed development for larvae cultured at 20—22°C. By contrast, when larvae of an unidentified, subtidal naticid were cultured at 12°C and fed Isochrysis galbana, they showed a slow but consistent rate of growth and development, and they achieved metamorphic competence at 3 to 3% months after hatching. Larvae of this unidentified species had a smooth protoconch I, un- like the sculptured protoconch I of P. lewisii, and they grew to a larger size at metamorphic competence than did veligers of P. lewisii (Pedersen, 1996). Our repeated failure to obtain a continuous progression of growth and development of P. lewisii larvae at 12°C, despite the fact that another species of naticid did well at this tempera- ture, encourages us to believe that our inability to rear P. lewisii at 12°C may not be a laboratory artifact. Summer seawater temperatures in the shallow, protected embay- ments inhabited by adults of P. lewisii typically rise well above 12°C during the late spring and summer when egg masses are abundant. Data for Saanich Inlet show that summer temperatures of 15—20°C are routine for surface waters and shallow embayments (Dr. Louis A. Hobson, University of Victoria, personal communication). We speculate that larvae of P. lewisii may generally stay within the relatively warm bays where they hatch, and in fact, they may need these warmer temperatures to com- plete larval development. Larvae of P. lewisii were induced to metamorphose by sediment from the intertidal zone of beaches inhabited by adults. The fact that sediment loses its inductive potency after autoclaving suggests that the active factor is organic, rather than some physical property of the sediment par- ticles. The relatively small percent metamorphosis in the presence of bivalves alone might have been due to active factor in residual sediment or organic surface film adher- ing to the bodies of the bivalves. Unfortunately, we did not specifically test the ability of ostracods to initiate metamorphosis of P. /ewisii. Until future experiments can narrow down the exogenous trigger for metamorphic in- duction, we tentatively propose that the active factor is R. V. K. Pedersen & L. R. Page, 2000 an indirect indicator of sites likely to harbor suitable small prey for P. lewisii. Finally, our results do not agree with Bernard’s (1967: 20) statement that young juveniles of P. lewisii feed on diatoms and Ulva until they are 5—6 mm in shell length, when they switch to predation on bivalves. We found that bivalves and ostracods were drilled and eaten by P. lew- isii within 3 to 5 days of metamorphic loss of the velum. We found no evidence of feeding on Ulva when this was added to bowls containing young juveniles. Our obser- vations are similar to those of Berg (1976), who reported that Natica gualtieriana is able to drill and ingest small gastropods shortly after metamorphosis. Kabat (1990) noted that the paleontological literature attributes bore- holes in fossil ostracod shells to naticid predation. He therefore speculated that ostracods may be a ‘“‘potentially important prey source for juvenile naticids.’’ Our obser- vations confirm that at least one species of extant naticid is an active predator of ostracods during its early post- metamorphic development. Acknowledgments. Supported by a research grant to L.R. Page from the Natural Sciences and Engineering Research Council of Canada. LITERATURE CITED Amio, M. 1955. 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Predation by the naticid gastropod Polinices lewisii (Gould) on little- neck clams Protothaca staminea (Conrad) in British Colum- bia. Canadian Journal of Zoology 72:319—325. Rew, R. G. B. & J. A. FRIESEN. 1980. The digestive system of the moon snail Polinices lewisii (Gould, 1847) with empha- sis on the role of the oesophageal gland. The Veliger 23:25— 34. RED, R. G. B. & B. D. GustaFson. 1989. Update on feeding and digestion in the moon snail Polinices lewisii (Gould, 1847). The Veliger 32:327. TuHorRSON, G. 1935. Studies on the egg-capsules and development of Arctic marine prosobranchs. Meddelelser om Grgnland 100:1-71. ZIEGELMEIER, E. VON. 1954. Beobachtungen tiber den Nahrung- serwerb bei der Naticidae Lunatia nitida Donovan (Gastrop- oda Prosobranchia). Helgolander Wissenschaftliche Meer- sesuntersuchungen 5:1—33. ZIEGELMEIER, E. VON. 1958. Zur Lokomotion bei Naticiden (Gas- tropoda Prosobranchia). Helgolander Wissenschaftliche Meersesuntersuchungen 6:202—206. The Veliger 43(1):64—71 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Polygyrid Land Snails, Vespericola (Gastropoda: Pulmonata), 3. Three New Species From Northern California BARRY ROTH AND WALTER B. MILLER Department of Invertebrate Zoology, Santa Barbara Museum of Natural History, Santa Barbara, California 93105, USA Abstract. Three new species of polygyrid land snails from northern California are described: Vespericola sasquatch, from stream drainages tributary to the Salmon River, Siskiyou County; Vespericola embertoni, from Reeves Canyon, Mendocino County; and Vespericola rhodophila, from coastal Sonoma County. INTRODUCTION This is the third in a series of studies by us on the sys- tematics of the western American polygyrid land snail genus Vespericola Pilsbry, 1939. In the first article of the series (Roth & Miller, 1993), we separated Vespericola pilosus (Henderson, 1928) from Vespericola columbianus (Lea, 1838), of which it had been considered a subspecies and showed that the distribution of V. pilosus is restricted to the San Francisco peninsula, California. We also dem- onstrated that Vespericola orius (Berry, 1933) is a spe- cies, not a subspecies of V. columbianus, and we de- scribed a new species, Vespericola marinensis Roth & Miller, 1993, from Marin County, California. In the second article of the series (Roth & Miller, 1995), we redescribed Vespericola megasoma (Pilsbry, 1928), adding details of its reproductive anatomy, and restricted its type locality to the east bank of Prairie Creek, near the south end of Prairie Creek Redwoods State Park, Humboldt County, California. We also de- scribed the reproductive anatomy of Vespericola eritri- chius (Berry, 1939) and Vespericola karokorum Tal- madge, 1962. Additionally, we described a new species, Vespericola klamathicus Roth & Miller, 1995, from drainages adjacent to those of V. karokorum in the Klam- ath Mountains, California. In a subsequent article (Cordero & Miller, 1995), one of us (WBM) assisted Alicia M. Cordero in describing the reproductive anatomy of Vespericola shasta (Berry, 1921), and in describing two new species, Vespericola rothi Cordero & Miller, 1995, and Vespericola scotti Cor- dero & Miller, 1995, from areas relatively close to but outside the distribution of V. shasta. In this third article by Roth & Miller we describe three new species. MATERIALS AnD METHODS Shell height and diameter are vernier caliper measure- ments and exclude the expanded lip of mature shells. Whorls were counted by the method of Pilsbry (1939:xi, fig. B). The density of periostracal setae was estimated by counting the number of setae per square millimeter on the shoulder of the body whorl, 0.25 whorl behind the aperture of adult specimens, at 30X magnification under a dissecting microscope with an ocular reticle. Three counts were taken per specimen and the mean (to the nearest integer) recorded. Specimens for dissection and whole mounts of geni- talia were prepared by the methods described by Roth & Miller (1993). The following abbreviations are used: ANSP, Academy of Natural Sciences of Philadelphia; BR, senior author’s collection, San Francisco, California; CAS, California Academy of Sciences; LACM, Los Angeles County Mu- seum of Natural History; SBMNH, Santa Barbara Mu- seum of Natural History. SYSTEMATICS POLYGYRIDAE Pilsbry, 1895 Vespericola Pilsbry, 1939 Vespericola Pilsbry, 1939:xvii. —Pilsbry, 1940:892-894. —Zilch, 1960:586. —Roth & Miller, 1993:135. Type species: Polygyra columbiana pilosa Henderson, 1928 [= Vespericola pilosus (Henderson)], by original designation. Vespericola sasquatch Roth & Miller, sp. nov. (Figures 1—5) Diagnosis: A large Vespericola with depressed-globose, almost imperforate shell, 5.8—6.25 whorls, erect, distant periostracal setae with forked bases, and usually a small parietal lamella. Penis medium-sized, slender, with ante- rior 50% enclosed in sheath; with short, conical, pointed verge 0.5—0.8 mm long; spermathecal duct massive, grad- ually tapering to a constriction at spermatheca. B. Roth & W. B. Miller, 2000 Page 65 Figures 1—3 Vespericola sasquatch Roth & Miller, sp. nov. Shell, holotype, SBMNH 143179, CALIFORNIA: Siskiyou County: along small stream entering Salmon River from north, 4.3 km E of Etna-Somesbar Road/California Highway 96 intersection at Somesbar (sec. 1 or 2, T. 11 N, R. 6 E, Humboldt Base and Meridian; USGS Forks of Salmon quadrangle [7.5-minute series, Topographic]). B. Roth, E. J. Kools coll., 17 May 1984. Top, apertural, and basal views. Diameter 16.0 mm. Figures 4, 5 Vespericola sasquatch Roth & Miller, sp. nov. Drawings made from projections of stained whole mounts. Structures seen in transparency are shown by dotted lines. Scale line = 1 mm. Figure 4. Anterior part of reproductive system, paratype, SBMNH 78125, CALIFORNIA: Siskiyou County: along small stream entering Salmon River from north, 4.3 km E of Etna-Somesbar Road/California Highway 96 intersection at Somesbar (sec. 1 or 2, T. 11 N, R. 6 E, Humboldt Base and Meridian; USGS Forks of Salmon quadrangle [7.5-minute series, Topographic]). B. Roth, W. B. Miller coll., 10 April 1992. Figure 5. Penis and penial sheath cut open to show verge and pilasters, paratype, SBMNH 77923, same locality as the preceding, B. Roth, E. J. Kools coll., 17 May 1984. Abbreviations: at, atrium; cp, cut edge of penis; cs, cut edge of penial sheath; ep, epiphallus; go, genital orifice; ov, oviduct; pe, penis; pi, pilaster; pr, penial retractor; ps, penial sheath; pt, prostate; re, retentor; sd, spermathecal duct; sp, spermatheca; ut, uterus; va, vagina; vd, vas deferens. Page 66 Description of shell: Shell (Figures 1—3) large for the genus, depressed-globose, almost imperforate, of 5.8— 6.25 whorls; base inflated. Spire low-domed, its sides weakly convex; whorls somewhat flattened, suture mod- erately impressed. Embryonic whorls 1.5—1.6; initial 0.2 whorl smooth; remainder of embryonic shell with crowd- ed, irregular, papillose, radiating rugae. Early teleoconch whorls with fine, slightly retractive growth rugae and dis- tant, erect or forwardly convex, acicular setae in convex- forward, protractive, descending rows. Most setae forked at base, furcae pointing aperturally; many setae with broad, finlike basal extension abaperturally. Periostracum between setae radially wrinkled, somewhat scaly on first four whorls, smoother on whorls five and six. Periphery rounded. Base regularly setose, setae smaller than on spire, extending into umbilical crevice. Last whorl not markedly descending, constricted behind lip. Aperture broadly auriculate, peristome concave in profile, oblique, at angle of about 40° to vertical; lip expanded and strong- ly reflected, moderately thickened submarginally, most strongly turned backward at base. Inner end of basal lip reflected over narrow, obliquely entering umbilical crev- ice. Parietal callus granulose, free edge convex, with only a very small re-entrant below upper limb of peristome. Small, white, straight parietal lamella usually present, set on upper third of parietal callus, outer edge on line be- tween upper and lower limbs of peristome. Shell tan, peri- stome pinkish tan. Dimensions of holotype: Diameter (exclusive of expand- ed lip) 16.0 mm, height 9.3 mm, whorls 6.25. Measurements and counts of material at hand: Range of adult shell diameter 13.5—16.0 mm (mean of five spec- imens including holotype, 15.0 mm); height 8.0—10.2 mm (x = 9.28 mm); height/diameter ratio 0.58-0.66 (xk = 0.618); number of whorls 5.8—6.25 (x = 6.13). Description of soft anatomy: The holotype and five paratypes were dissected. Living animal pinkish buff, darker and grayer on body- stalk. Mantle over lung clear buff, 25—40% maculated with black. Atrium (Figure 4) of moderate length for the genus. Penis elongate-conical, with anterior, basal half enclosed in diaphanous sheath adnate to base. Penial retractor mus- cle inserted on epiphallus. Narrow retentor muscle ex- tending from penial retractor muscle at attachment on epi- phallus to summit of penial sheath, from which other thin retentor fibers connect with parts of epiphallus and vas deferens. Sheathed part of penis about 4.0 mm long; pro- truding part 2.8-4.5 mm long. Short peduncular section of about 0.5 mm present between base of sheath and junc- tion with atrium. Apex of penis containing short, conical, pointed verge, 0.5—0.8 mm long, through which seminal duct opens into penial chamber (Figure 5). Spermathecal duct massive, tightly appressed to free The Veliger, Vol. 43, No. 1 oviduct (which is smaller in diameter and branches from it), with one deep convolution below spermatheca, cylin- drical-conic, about 6.0 mm long, about 2.0 mm in di- ameter at junction with oviduct, gradually tapering to 1.0 mm constriction at base of spermatheca. Spermatheca varying from oblong-ovate to teardrop-shaped, about 4.0 mm long, with bluntly pointed tip. Type material: Holotype: SBMNH 143179 (shell and dissected anatomy), CALIFORNIA: Siskiyou County: along small stream entering Salmon River from north, 4.3 km E of Etna-Somesbar Road/California Highway 96 in- tersection at Somesbar (sec. 1 or 2, T. 11 N, R. 6 E, Humboldt Base and Meridian; USGS Forks of Salmon quadrangle [7.5-minute series, Topographic]). B. Roth, E. J. Kools coll., 17 May 1984. Paratypes: SBMNH 77923 (2), from same locality as holotype; SBMNH 78125 (3) from same locality as ho- lotype, B. Roth, W. B. Miller coll., 10 April 1992. Ad- ditional paratypes, ANSP, BR and CAS. Referred material: CALIFORNIA: Siskiyou County: Scott River Road, W of Hotelling Gulch. M. K. Gausen coll., November 1998 (BR 2133); near lower Salmon River, NE1/4 sec. 2, T. 11 N, R. 6 E, Humboldt Base and Meridian. T. Hacking coll., 22 June 1999 (BR 2230). Remarks: Vespericola sasquatch is distinguished from most other species in northwestern California by its large size, large number of whorls, and depressed-globose shape. The combination of large shell and small parietal tooth is unique to V. sasquatch, but in one of the adult specimens examined the parietal tooth is absent. Vesper- icola karokorum Talmadge, 1962, from streams draining into the Klamath River near Orleans, Humboldt County, differs in having fewer, more distantly spaced, and longer periostracal setae, many of which have recurved tips. The embryonic whorls of V. karokorum are more coarsely pa- pillose. The inner lip is reflected over the umbilicus to about the same degree in both species; but the base of V. sasquatch is somewhat more flattened, that of V. karo- korum more inflated. Vespericola karokorum has an acic- ular verge, about 3.4 mm long; the verge of V. sasquatch is short and conical, 0.5—0.8 mm long. The vegetation at the type locality is mixed woodland with Douglas fir (Pseudotsuga menziesii), canyon oak (Quercus chrysolepis), bigleaf maple (Acer macrophyl- lum), and madrone (Arbutus menziesii). Vespericola sas- quatch was found under logs on the ground, particularly near the stream bed. Other mollusks found with V. sas- quatch at the type locality included Prophysaon ander- soni (Cooper, 1872), Monadenia fidelis salmonensis Tal- madge, 1954, and Ancotrema sp., cf. A. voyanum (New- comb, 1865). Etymology: Sasquatch, from the Salish language: a hairy, anthropoid being said to inhabit northwestern B. Roth & W. B. Miller, 2000 Page 67 Figures 6-8 Vespericola embertoni Roth & Miller, sp. nov. Shell, holotype, SBMNH 143178, CALIFORNIA: Mendocino Coun- ty: tributary ravine to Reeves Canyon, on W side of U.S. Highway 101 at milepost 54.60 (3.9 road mi N of Outlet Creek bridge), W. B. Miller coll., 7 April 1990. Top, apertural, and basal views. Diameter 15.4 mm. North America, including the Klamath Mountains. For purposes of the American Fisheries Society list of the common names of mollusks (Turgeon et al., 1998) and other administrative uses, we propose the name “‘sas- quatch hesperian.”’ Vespericola embertoni Roth & Miller, sp. nov. (Figures 6—10) Diagnosis: A large Vespericola with depressed-globose, very narrowly umbilicate shell, 5.4—6.1 whorls, sparse periostracal setae, and sometimes a small parietal lamella. Penis elongate-conical, with anterior 60% of length en- closed in sheath; with short, conical, pointed verge 1.0 mm long; spermathecal duct massive, swelling to a di- ameter of 2.0 mm before tapering to a constriction at base of spermatheca. Description: Shell (Figures 6—8) large for the genus, de- pressed-globose, very narrowly umbilicate, of 5.4—6.1 whorls; base inflated. Spire low-domed to broadly coni- cal, its sides straight or weakly convex; whorls flattened, suture moderately impressed. Embryonic whorls 1.4—1.8; initial 0.1 whorl smooth; remainder of embryonic shell with crowded, irregular, papillose, radiating rugae. Post- embryonic sculpture of low, retractive rugae and dense, sharp granulation with collabral trend. Periostracum bear- ing sparse, moderately long, acicular setae in distant, steeply descending rows; 3—4 setae/mm? on shoulder of body whorl, erect or curving away from direction of coil- ing, occasionally forked at base, usually with finlike ab- apertural basal extension. Periostracum between setae sharply granulose and finely wrinkled, with a few fine, raised spiral lirae on shoulder of whorl. Periphery round- ed, sometimes more sharply curved at shoulder. Base tu- mid, densely setose with setae shorter than on spire, pap- illose where setae worn off. Last whorl not markedly de- scending, constricted behind lip. Aperture broadly auric- ulate; peristome concave in profile, oblique, at angle of 35—45° to shell axis; lip expanded and reflected, lightly to moderately thickened submarginally. Inner part of bas- al lip narrowed, weakly deflected forward, end dilated backward over narrow umbilicus. Parietal callus granu- lose, free edge convex, with shallow re-entrant below up- per limb of peristome. Small, white, convex-forward pa- rietal lamella present in about two-thirds of all adult spec- imens examined. Shell tan; lip pinkish tan. Dimensions of holotype: Diameter (exclusive of expand- ed lip) 15.4 mm, height 9.5 mm, whorls 6.0. Measurements and counts of material at hand: Range of adult shell diameter 13.2—16.6 mm (mean of 25 spec- imens including holotype, 14.95 mm); height 8.0—10.0 mm (x = 9.14 mm); height/diameter ratio 0.57—0.66 (x = 0.611); number of whorls 5.4—6.1 (K = 5.85). Description of soft anatomy: The holotype and eight paratypes were dissected. Color of living animal pinkish buff, darker and grayer on body-stalk. Mantle over lung clear buff, 20—40% mac- ulated with black. Atrium (Figure 9) of moderate length for the genus. Penis elongate-conical, with anterior, basal portion en- closed in thin sheath adnate to base. Penial retractor mus- cle inserted on epiphallus. Narrow retentor muscle ex- tending from penial retractor muscle at attachment on epi- phallus to summit of penial sheath, from which other thin retentor fibers form connections with parts of epiphallus and vas deferens. Sheathed part of penis in holotype about 5.5 mm long; protruding part about 3.6 mm. In mature paratypes, sheathed part varying from 5.0 to 6.2 mm, with a mean of 5.6 mm; protruding part varying from 3.0 to 4.8 mm, with a mean of 4.1 mm. Mean ratio of protruding length to sheathed length about 0.8. Slender Page 68 The Veliger, Vol. 43, No. 1 Figures 9, 10 Vespericola embertoni Roth & Miller, sp. nov. Drawings made from projections of stained whole mounts. Structures seen in transparency are shown by dotted lines. Scale line = 1 mm. Figure 9. Anterior part of reproductive system of paratype, SBMNH 77888, CALIFORNIA: Mendocino County: tributary ravine to Reeves Canyon on W side of U.S. Highway 101 at milepost 54.60 (3.9 road mi N of Outlet Creek bridge), B. Roth, W. B. Miller coll., 19 November 1989. Figure 10. Penial complex of paratype, SBMNH 77888, with apical portion of penis opened to show verge and pilasters; same locality as the preceding, W. B. Miller coll., 7 April 1990. peduncular section of about 1.4 mm present between base of sheath and junction with atrium. Apex of penis con- taining short, conical, pointed verge 1.0 mm long and 0.4 mm wide at base (Figure 10). Seminal duct opening into penial chamber at tip of verge. Spermathecal duct massive, tightly appressed to free oviduct (which is smaller in diameter and branches from it), cylindrical-conic, about 5.5 mm long, about 1.4 mm in diameter at junction with oviduct, widening to a max- imum diameter of 2.0 mm before tapering to a 0.5 mm constriction at base of spermatheca. Spermatheca oblong- ovate in fully mature specimens, often flattened on side apposed to prostate/uterus and narrowly cylindrical in less mature individuals, about 5.5 mm long, with rounded tip. Type material: Holotype: SBMNH 143178 (shell and dissected anatomy), CALIFORNIA: Mendocino County: tributary ravine to Reeves Canyon, N of Willits, on W side of U.S. Highway 101 at milepost 54.60 (3.9 road mi N of Outlet Creek bridge), W. B. Miller coll., 7 April 1990. Paratypes: SBMNH 77888 (8), from same locality as holotype. Additional paratypes, ANSP, BR, CAS. B. Roth & W. B. Miller, 2000 Page 69 Figures 11-13 Vespericola rhodophila Roth & Miller, sp. nov. Shell, holotype, SBMNH 000000, CALIFORNIA: Sonoma County: China Gulch at crossing of Kruse Ranch Road, 1.1 road km from California Highway 1, Kruse Rhododendron State Reserve. W. B. Miller, B. Roth coll., 17 November 1989. Top, apertural, and basal views. Diameter 13.2 mm. Referred material: CALIFORNIA: Humboldt County: Richardson Grove State Park (BR 1816). Mendocino County: just south of Longvale (CAS 052365); Reeves Canyon S of Longvale along US Highway 101 at mile- post 53.91, in redwood stump (BR 336); Reeves Canyon, in ravines off US Highway 101, 7.8 mi N of Willits, at mileposts 54.60 and 53.94 (SBMNH 77999); Reeves Canyon, on W side of US Highway 101, 7.8 mi N of Willits, at milepost 54.11, along tributary ravine, under logs and in old stumps (SBMNH 77866). Remarks: A parietal lamella is variably present or ab- sent. The single specimen from Richardson Grove, Hum- boldt County, lacks a lamella. It has 2—3 periostracal se- tae/mm?’, compared to 3—4 setae/mm* in the type lot. Vespericola embertoni is distinguished anatomically from other species by its short, conical, pointed verge at the apex of a slender, elongate-conical penis sheathed ba- sally for about half to two-thirds of its length, and by a massive spermatheca and spermathecal duct complex as large or larger than the penial complex. The large, depressed shell, sparse setation, very narrow umbilicus, and granular surface texture distinguish V. em- bertoni from most other species in northern California. The type lot was collected in a stream gully in second- growth redwood forest. The predominant tree species along the riparian corridor was tan oak (Lithocarpus den- siflora), with Douglas fir (Pseudotsuga menziesii) and California bay (Umbellularia californica) also prominent. Other land mollusks found at the type locality included Haplotrema minimum (Ancey, 1888), Monadenia infu- mata (Gould, 1855), and Helminthoglypta arrosa pom- oensis A. G. Smith, 1938. Etymology: The species is named for Kenneth C. Em- berton, in honor of his innovative research on the family Polygyridae. For purposes of the American Fisheries So- ciety list of the common names of mollusks (Turgeon et al., 1998) and other administrative uses, we propose the name “‘Reeves Canyon hesperian.”’ Vespericola rhodophila Roth & Miller, sp. nov. (Figures 11—16) Diagnosis: A medium-sized Vespericola with depressed- globose, almost imperforate shell of 5.5—5.8 whorls; peri- ostracal setae about 30—36/mm~°. Penis either completely enclosed by sheath or with upper 10—20% swollen and protruding; with 1.5 mm, cylindrical-conic verge with membranous, lateral flaps and tubular ventral groove forming outlet of seminal duct. Description: Shell of medium size for the genus, de- pressed-globose, almost imperforate, of 5.5—5.8 whorls; base inflated, solid-looking. Spire broadly conic, its sides weakly convex; whorls somewhat flattened, suture shal- lowly impressed. Embryonic whorls 1.6, sculptured with crowded, radially elongate, blunt papillae that on the sec- ond whorl tend to align in radiating rows separated by shallow grooves. Early neanic whorls with fine, slightly retractive growth rugae and rather sparse, erect or gently curving, acicular setae in protractive, descending rows. Setae not obviously forked at base; some setae with fin- like basal extension abaperturally. Periostracum between setae radially wrinkled and irregularly granulose. Setae closer together, shorter, and more regularly spaced on subsequent whorls, 30—36/mm/ on body whorl. Periphery rounded, broadest above middle of whorl, somewhat sloping toward base. Base densely and regularly setose. Last whorl not markedly descending, sharply constricted behind lip. Aperture broadly auriculate, peristome con- cave in profile, oblique, at angle of about 45° to vertical; lip expanded and reflected, most strongly at base, mod- erately thickened submarginally. Umbilical crevice ex- tremely narrow, oblique. Basal lip straight; inner lip nar- Page 70 The Veliger, Vol. 43, No. 1 Figures 14—16 Vespericola rhodophila Roth & Miller, sp. nov. Drawings made from projection of stained whole mounts. Scale line = 1 mm. Figure 14. Anterior portion of reproductive system of holotype, SBMNH 00000, CALIFORNIA: Sonoma County: China Gulch at crossing of Kruse Ranch Road, 1.1 road km from California Highway 1, Kruse Rhododendron State Reserve. W. B. Miller, rowed, weakly curved forward, and dilated so that it en- croaches on, and nearly covers, umbilicus from left side. Parietal callus granulose, free edge strongly convex, swinging well to left of umbilicus, with shallow sinus below upper limb of peristome. Moderate-sized, white, straight parietal lamella usually present near middle of parietal callus. Shell light reddish brown; peristome pink- ish tan, with whitish callus thickening. Dimensions of holotype: Diameter (exclusive of expand- ed lip) 13.2 mm, height 9.0 mm, whorls 5.5. Soft anatomy: The color of living animals is pinkish buff, darker and grayer on the body-stalk. The mantle over the lung is clear buff, 20—40% maculated with black. The holotype and 13 paratypes were dissected; an ad- ditional five specimens from Salt Point State Park were also dissected. The atrium (Figure 14) is of moderate length for the genus. The penis is swollen, banana-shaped, with a bulbous apex, its anterior, basal portion enclosed in a thin sheath adnate to the base, and the apical protruding part of wider diameter than the sheathed part. The penial retractor mus- cle is inserted on the epiphallus. A narrow retentor mus- cle extends from the penial retractor muscle at its attach- ment on the epiphallus to the summit of the penial sheath, from which other thin retentor fibers form connections with parts of the epiphallus and vas deferens. The sheathed part of the penis in the holotype is about 6.0 mm long; the protruding part is about 2.5 mm. In the mature paratypes, the sheathed part varies from 4.4 to 6.5 mm, with a mean of 5.8 mm; the protruding part varies from 2.0 to 5.0 mm, with a mean of 3.3 mm. The mean ratio of protruding length to sheathed length is about 0.6. There is a slender peduncular section of about 1.5 mm between the base of the sheath and the junction with the atrium. The apex of the penis contains a short, tongue-shaped, verge 1.0 mm long and 0.2 mm wide at its base, equipped for the apical two-thirds of its length with a narrow, ven- tral, longitudinal groove flanked by a thin membranous flap on each side, ending at the apex with an open groove about 0.4 mm long, which forms the outlet of the seminal duct (Figure 15). The inner wall of the penis consists of a reticular pattern of closely appressed papillae (Figure 16). The spermathecal duct is massive and short, appressed <— B. Roth coll., 17 November 1989. Figure 15. Lateral, ventral, and dorsal views of verge of paratype SBMNH 77951. Figure 16. Anterior portion of reproductive system of paratype, SBMNH 77951, with apical portion of penis opened to show verge and internal papillae; same locality as the preceding, W. B. Miller, B. Roth coll., 6 October 1990. B. Roth & W. B. Miller, 2000 Page 71 to the free oviduct (which is smaller in diameter and branches from it); it is cylindrical-conic, about 3.0 mm long, with a diameter of about 1.5 mm at its junction with the oviduct, tapering gradually to a 0.5 mm constriction at the base of the spermatheca. The spermatheca is oblong-ovate in fully mature spec- imens and narrowly cylindrical in less mature individuals, about 3.3 mm long, with a rounded tip. Type material: Holotype: SBMNH 00000 (shell and stained whole mount of reproductive system), CALIFOR- NIA: Sonoma County: China Gulch at crossing of Kruse Ranch Road, 1.1 road km from California Highway 1, Kruse Rhododendron State Reserve. W. B. Miller, B. Roth coll., 17 November 1989. Paratypes: SBMNH 77860 (4), from same locality as holotype. SBMNH 77951 (10), along China Gulch creek at crossing of Plantation Road, 0.9 km from California Highway 1, Kruse Rhododendron State Reserve. W. B. Miller, B. Roth coll., 6 October 1990. Additional para- types, ANSP, BR, CAS. Referred material: CALIFORNIA: Sonoma County: Kruse Rhododendron State Reserve, below parking lot (SBMNH 75030); Miller Creek, at crossing of California Hwy. 1 (first deep hairpin turn N of Gerstle Cove Camp- ground), Salt Point State Park (BR 1672, BR 1712, WBM 7861). Remarks: In the material at hand, adult shell diameter ranges from 11.9 to 13.7 mm (mean of 25 specimens including holotype, 13.02 mm); height, 8.2 to 9.9 mm (x = 8.96 mm); height/diameter ratio, 0.64 to 0.76 (xk = 0.688); number of whorls, 5.2 to 5.8 (x = 5.54). In some specimens, the body whorl is pale straw-col- ored rather than reddish brown. Vespericola rhodophila is distinguished anatomically from other species by its banana-shaped penis with a bul- bous apical part usually protruding from the basal, sheathed part. The short, grooved verge with membranous lateral flaps somewhat recalls the spoon-shaped verge of V. megasoma but is not as stout. The species is found under logs in riparian woodland. Etymology: Gr., rhodos, rose, + philos, lover, with ref- erence to the Western Azalea (Rhododendron occidenta- le) which is a prominent component of the understory within its range. For purposes of the American Fisheries Society list of the common names of mollusks (Turgeon et al., 1998) and other administrative uses, we propose the name “‘azalea hesperian.”’ Acknowledgments. We are grateful to Elizabeth J. Kools for help in the field during the collection of the original material of Vespericola sasquatch and to Mary K. Gausen and Tony Hack- ing for specimens from additional localities. LITERATURE CITED CorRDERO, A. M. & W. B. MILLER. 1995. Reproductive anatomy of Vespericola shasta (Berry, 1921) (Gastropoda: Pulmona- ta: Polygyridae), and descriptions of two new species of Ves- pericola from northern California. The Veliger 38:304—311. HENDERSON, J. 1928. Polygyra columbiana pilosa, new subspe- cies. The Nautilus 41(4):143. PitsBry, H. A. 1939. Land Mollusca of North America (north of Mexico). Academy of Natural Sciences of Philadelphia Monograph 3, 1(1):i—xvii, 1-573, i-ix. Pitssry, H. A. 1940. Land Mollusca of North America (north of Mexico). Academy of Natural Sciences of Philadelphia Monograph 3, 1(2):i-—vil, 574—994, i-ix. Rotnu, B. & W. B. MILLER. 1993. Polygyrid land snails, Vesper- icola (Gastropoda: Pulmonata), 1. Species and populations formerly referred to Vespericola columbianus (Lea) in Cal- ifornia. The Veliger 36:134—144. Rot, B. & W. B. MILLER. 1995. Polygyrid land snails, Vesper- icola (Gastropoda: Pulmonata), 2. Taxonomic status of Ves- pericola megasoma (Pilsbry) and V. karokorum Talmadge. The Veliger 38:133-144. TALMADGE, R. R. 1962. A new land snail from the Klamath Mountains, California (Mollusca: Pulmonata: Polygyridae). The Veliger 5:28—29, pl. 5. TuRGEON, D. D., J. EF QuINN Jr., A. E. BoGAN, E. V. COAN, E G. HOcHBERG JR., W. G. Lyons, P. M. MIKKELSEN, R. J. NEVES, C. E E. Roper, G. ROSENBERG, B. ROTH, A. SCHELTEMA, F. G. THompson, M. VECCHIONE & J. D. WILLIAMS. 1998. Com- mon and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. 2nd ed. American Fisheries Society Special Publication 26:i—x, 1-526. ZiLcu, A. 1959-1960. Gastropoda, Teil 2, Euthyneura. Handbuch der Paléiozoologie 6(2):1—400 (1959); 401-834 (1960). The Veliger 43(1):72—77 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Distribution of the Bonnet Limpet, Hipponix conicus (Gastropoda: Hipponicidae), among Host Species in Western Kyushu, Japan KAZUNORI YAMAHIRA' AnD FUMITO YANO Amakusa Marine Biological Laboratory, Kyushu University, Tomioka, Reihoku-cho, Amakusa, Kumamoto 863-25, Japan Abstract. The bonnet limpet, Hipponix conicus (Schumacher, 1817), adheres to the shell surface of other snail species. In order to discuss the ecological relationships between H. conicus and many host snails, the distribution of this limpet among its potential hosts was determined in western Kyushu, Japan. A total of 26 snail species was collected, and H. conicus was found on 13 of them. The mean adhesion frequencies and mean numbers of H. conicus per snail were significantly different among the host species. Among eight host species, more H. conicus were found on shells with living snails than on shells with hermit crabs. The number of H. conicus per host and shell length of H. conicus tended to increase with host size for 11 and eight host species, respectively. Host snail individuals with at least one H. conicus tended to be larger than those without any H. conicus for ten species. Overall, our data indicate that the quality of shells as a host differs among host species (living snails and hermit crabs) or host sizes. The difference in H. conicus loads could be caused primarily by preferential adhesion to different hosts and/or differential growth and/or survival on different hosts. INTRODUCTION The bonnet limpet, Hipponix conicus (Schumacher, 1817), attaches to shells of many different species of ma- rine gastropods in Japan, including commercially impor- tant ones, such as the Japanese abalone, Haliotis aquatilis (Reeve, 1846) and the turban shell, Turbo cornutus (Lightfoot, 1786) (Matsunaga, 1964). However, bivalves never harbor H. conicus (Matsunaga, 1964). Hipponix conicus reproduces on host snails by direct development (i.e., there is no pelagic stage) (Habe, 1953; Amio, 1963); and several authors (Yonge, 1953, 1960; Cernohorsky, 1968; Knudsen, 1991) postulated that it is a protandrous, consecutive hermaphrodite. Hipponix conicus is often observed to deeply erode the shell of its host species (Knudsen, 1991) and therefore has been suggested to reduce their market value (Mat- sunaga, 1964). In particular, it appears that H. conicus adheres more strongly to cultured varieties of host species than to natural forms, probably resulting in severe eco- nomic damage. To date, however, little attention has been paid to interactions between H. conicus and host species. Matsunaga (1964) investigated the ecological relationship between H. conicus and Haliotis aquatilis with respect to the adhesional position on the host and movement be- tween host individuals. He stated that H. conicus tended Present address: Department of Environmental Chemistry, Fac- ulty of Engineering, Kyushu Kyoritsu University, Kitakyushu 807-8585, Japan; e-mail: yamahira@kyuko-u.ac.jp to adhere near respiratory pores to eat feces or soft parts of the host, and that H. conicus sometimes moved among host individuals in pursuit of better habitats. On the other hand, Knudsen (1991) argued that it is unlikely that fecal pellets of the host would constitute an essential part of the food of Hipponix australis (Lamarck, 1819), similar to and taxonomically confused with H. conicus, and that H. australis is not selective as to its choice of gastropod host, because a wide variety is used. However, the earlier authors did not conduct a quantitative survey of the pref- erence of Hipponix species for host gastropod species, which would be primary information regarding the eco- logical relationship among them. This paper describes the distribution of H. conicus among many potential host species inhabiting intertidal to shallow subtidal zones, and examines some ecological relationships between H. conicus and the host species. In particular, the differential H. conicus loads among differ- ent species and different sizes of hosts are discussed from the viewpoint of preferential adhesion to different hosts (host preference), differential growth and/or survival on different hosts (host quality), and differences in exposure to H. conicus among hosts (opportunity for adhesion). MATERIALS AND METHODS Our study was carried out during the spring low tides of October 1994 at Magarisaki spit, Amakusa Shimo-shima Island, western Kyushu, Japan (32°32'N, 130°2’E). Fifty- six 50 X 50 cm quadrats were set in the low intertidal K. Yamahira & F. Yano, 2000 (0.4 m above mean low water spring tide: MLWS) and subtidal zone (0.6—2.6 m below MLWS). One or two lay- ers of boulders (mean diameter approx. 20—30 cm) cov- ered the substrate in all quadrats. On the beach, grazer snails and chitons are abundant (Takada & Kikuchi, 1990). All living snails and shells with hermit crabs in each quadrat were collected, transported to the laboratory, and fixed in 10% neutralized formalin. SCUBA diving was used for collecting in the subtidal zone. In the lab- oratory, all snails and shells were identified, and their size was measured with vernier calipers (+0.05 mm). Either shell length or shell width was used as a size indicator of each of the snail species. Because shell length repre- sents their sizes better than shell width for some species and vice versa for the others, we used appropriate mea- surements for each species. All shells were examined for the presence of Hipponix conicus. Adhesion frequencies of each host species (proportion of individuals with at least one H. conicus) were calculated for each quadrat, and the number of H. conicus on each shell was counted; then the shell length (length between the anterior and pos- terior edge) of each H. conicus was measured with the vernier calipers. Almost all of the H. conicus individuals remained on the shells through the fixation and handling (only a few were observed off from their hosts). Thus, the observed distribution patterns of H. conicus among hosts were considered to be natural, not artifactual. Kruskal-Wallis tests were used to determine if there were significant differences among host species in the adhesion frequencies in each quadrat and in the number of individuals per host. To compare H. conicus adhesion frequencies and numbers on shells with living snails and those with hermit crabs, Mann-Whitney U tests were used for each of the nine snail species whose shells were uti- lized by both H. conicus and hermit crabs. For each of the 12 snails species that harbor H. conicus, we deter- mined if the number and size of H. conicus was signifi- cantly correlated with the size of its host. Furthermore, to determine if larger hosts were more likely to harbor H. conicus, we compared the shell sizes of snails with and without H. conicus for each host species using Mann- Whitney U tests. RESULTS A total 26 snail species were collected from the site (Ta- ble 1); six species (23%) were found only in the low intertidal zone, 13 species (50%) were only in the subtidal zone, and seven species (27%) were found in both. Thir- teen species (50%) had at least one Hipponix conicus at- tached. Hipponix conicus was found on snail species col- lected from both tidal zones. However, only two Tectus pyramis individuals were collected, and they are excluded in all analyses below. No H. conicus was found living attached on the surface of boulders. Mean adhesion frequencies ranged from 0.04 (Japeu- Page 73 thria ferrea) to 1.0 (Thais bronni) (Table 2), and they were significantly different among host species (Kruskal- Wallis test, P < 0.0001). Mean numbers of H. conicus per host individual ranged from 0.08 (J. ferrea) to 6.17 (T. bronni) (Table 3) and were significantly different among host species (Kruskal-Wallis test, P < 0.0001). In host species that have higher adhesion frequencies, indi- vidual snails tended to carry more H. conicus: there was a significant positive correlation between the mean ad- hesion frequencies for a species and mean number of H. conicus per individual (r = 0.86, P = 0.0003, n = 12). The mean adhesion frequencies were higher on living snails than on hermit crab shells for all nine species (Ta- ble 2). The adhesion frequencies were significantly dif- ferent for five of the nine species, though the sample size of hermit crabs was rather small. The mean numbers of H. conicus per shell were also higher on living snails than on hermit crab shells for eight of the nine species (Table 3), and the differences were significant in five of the eight species. In general, the number of H. conicus per host increased with host size (Table 4). In five of the 12 species, there was significant positive correlation between the number of H. conicus per host and host size. H. conicus shell length also increased with host size in eight of the 11 host species (Table 5), but the correlation was significant in only two of the eight species. Furthermore, snaii in- dividuals with at least one H. conicus were larger than snails without any H. conicus in ten of the 11 host species (Table 6), and in three of the ten species, the difference was significant. DISCUSSION Although we observed that Hipponix conicus loads dif- fered among host species, these differences could be caused by preferential adhesion to different host species (host preference), differential growth and/or survival on different host species (host quality), or differences in ex- posure to H. conicus among host species. Comparison between the snail species frequently adhered to and those infrequently or never adhered to appears to say little about the conditions under which H. conicus adheres. No predictions about a species’ H. conicus load can be made based on taxonomic group, morphological features, or foraging habits. However, it seems that H. conicus is in- frequently found on species with the following charac- teristics: (1) small size (e.g., Mitrella bicincta, Trochus sacellus), (2) ability to cover themselves with a mantle (e.g., Purpuradusta gracilis, Adusta onyx); (3) covered by calcareous algae (e.g., Astralium haemafragum); and (4) ability to move to the higher intertidal zone (Mono- donta labio, Japeuthria ferrea; see Takada, 1996; N. Ota, unpublished data). These characteristics might decrease the shell’s quality as a host or the exposure time to H. conicus. Page 74 The Veliger, Vol. 43, No. 1 Table 1 Mean shell size (+SD) of living snails collected at the study site. Either shell length (SL) or shell width (SW) was measured as a size indicator of each of the snail species. Y (Yes) and N (No) indicate whether the species has at least one Hipponix conicus attached or not, respectively. Presence of Hipponix Snail species Measured part Mean + SD mm (n) conicus Found only in the intertidal zone Clanculus ater (Pilsbry, 1901) SW 7.18 + 5.20 (2) N Monodonta labio (Tapparone-Canefri, 1874) SW ' 13.64 + 1.89 (31) N Chlorostoma argyrostoma (Tapparone-Canefri, 1874) SW 21.23 + 2.91 (67) Y Lunella coronata (Récluz, 1853) SW 19.70 + 1.28 (132) SC Japeuthria ferrea (Reeve, 1847) SL 22.66 + 4.80 (12) Y Thais clavigera (Kiister, 1858) SL 24.40 (1) N Found in both the intertidal and subtidal zone Cantharidus japonicus (Adams, 1853) SL 6.77 + 1.59 (GB) N Omphalius nigerrima (Gmelin, 17919) SW 18.33 + 1.25 (10) Y Omphalius rusticus (Philippi, 1846) SW 17.55 + 4.42 (181) NC Turbo stenogyrum (Fischer, 1873) SW 16.26 + 5.24 (7) Ys Ergalatax contractus (Reeve, 1846) SL 17.75 + 3.92 (367) Y: Mitrella bicincta (Gould, 1860) SL 9.44 + 1.11 (26) N Pyrene scripta (Lamarck, 1822) SL 14.28 + 1.66 (182) WG Found only in the subtidal zone Calliostoma unicum (Dunker, 1860) SW 16.93 + 2.98 (3) N Omphalius pfeifferi (Philippi, 1846) SW 21.15 + 3.19 (40) BY, Trochus sacellus (Dunker, 1862) SW 16.10 (1) N Tectus pyramis (Born, 1778) SW 26.80 + 6.15 (2) NY? Astralium haemafragum (Menke, 1829) SW 18.49 + 3.92 (49) YC Conomurex luchuanus (Linnaeus, 1758) SL 58.15 (1) N Adusta onyx (Linnaeus, 1758) SL 39.85 (1) N Purpuradusta gracilis (Schilder, 1931) SL 15.95 + 1.78 (6) N Thais echinata (Blainville, 1832) SL 40.25 (1) N Thais bronni (Dunker, 1860) SL 27.43 + 10.11 (6) Nya Thais luteostoma (Holten, 1803) SL 24.33 + 6.11 (42) Y Mitra scutulata (Gmelin, 1791) SL 28.65 (1) N Pusia inermis (Reeve, 1845) SL 10.70 (1) N Table 2 Mean adhesion frequency (+SD) of each host shell with living snails and hermit crabs. Mann-Whitney U test: * and ** denote the significant difference at 5% and 1% level, respectively. Mean + SD (n) Host species Living snail Hermit crab P Thais bronni 1.00 + 0.00 (5) — Omphalius nigerrima 0.96 + 0.09 (5) 0.00 + 0.00 (2) 0.0134* Thais luteostoma 0.95 + 0.14 (23) 0.00 + 0.00 (2) 0.001 ** Omphalius pfeifferi 0.94 + 0.14 (19) 0.75 + 0.50 (4) 0.5394 Omphalius rusticus 0.63 + 0.32 (25) 0.11 + 0.33 (9) 0.0004** Chlorostoma argyrostoma 0.38 + 0.32 (13) 0.10 + 0.32 (10) 0.018* Lunella coronata 0.38 + 0.17 (16) 0.25 + 0.50 (4) 0.1293 Ergalatax contractus 0.32 + 0.29 (51) 0.06 + 0.25 (16) 0.0003** Astralium haemafragum 0.27 + 0.38 (25) — Turbo stenogyrum 0.13 + 0.25 (4) 0.00 (1) 0.6171 Pyrene scripta 0.09 + 0.18 (46) 0.00 + 0.00 (3) 0.2501 Japeuthria ferrea 0.04 + 0.12 (8) — K. Yamahira & F. Yano, 2000 Page 75 Table 3 Mean number (+SD) of Hipponix conicus per host shell with living snails and hermit crabs. Mann-Whitney U Test: * and ** denote the significant difference at 5% and 1% level, respectively. Mean + SD (n) Host species Living snail Thais bronni 6.17 + 4.58 (6) Thais luteostoma 5.88 + 5.64 (42) Omphalius pfeifferi 4.35 + 3.89 (40) Omphalius nigerrima 1.90 + 1.20 (10) Turbo stenogyrum esj7/ a= S05) (@) Omphalius rusticus 1.43 + 2.02 (181) Ergalatax contractus 0.84 + 1.34 (367) Chlorostoma argyrostoma 0.78 += 1.00 (67) Lunella coronata 0.61 + 0.98 (132) Astralium haemafragum 0.39 + 1.04 (49) Pyrene scripta 0.13 + 0.48 (182) Japeuthria ferrea 0.08 + 0.29 (12) Hermit crab P 0.00 + 0.00 (2) 0.0271* 5.50 + 6.14 (4) 1.0000 0.00 + 0.00 (2) 0.0478* 0.00 (1) 0.5677 0.04 + 0.20 (24) 0.0001 ** 0.18 + 0.85 (22) 0.0013** 0.05 + 0.23 (19) 0.0012** 0.33 + 0.82 (6) 0.4037 0.00 + 0.00 (3) 0.5677 Hipponix conicus was more likely to be found on living snails than on shells inhabited by hermit crabs. This fact suggests that living snails might promote growth and/or survival of H. conicus. Matsunaga (1964) reported that H. conicus uses its prolonged proboscis to eat the feces and mantle of the host snail. Laws (1971) concluded that H. conicus is a particle feeder that benefits from the par- ticles carried with the afferent current produced by hosts. On the other hand, Knudsen (1991) argued that fecal pel- lets of hosts do not constitute an essential part of the food of Hipponix. Larger hosts possessed more, larger H. conicus, prob- ably due to preferential adhesion to larger hosts (host preference), differential growth, and/or survival on dif- ferent-sized hosts (host quality). Large hosts may provide Table 4 Correlation coefficients (7) between host size and number of Hipponix conicus per host. Fisher F to z test: * and ** indicate that the correlation was statistically significant at 5% and 1% level, respectively. Host species r (n) P Thais bronni 0.080 (6) 0.8893 Thais luteostoma 0.543 (42) 0.0001 ** Omphalius pfeifferi 0.476 (40) 0.0017** Omphalius nigerrima —0.374 (10) 0.2990 Omphalius rusticus 0.509 (181) <0.0001** Chlorostoma argyrostoma 0.233 (67) 0.0571 Lunella coronata 0.183 (132) 0.0359* Astralium haemafragum 0.131 (49) 0.3710 Ergalatax contractus 0.258 (367) <0.0001** Turbo stenogyrum 0.674 (7) 0.1021 Pyrene scripta 0.127 (182) 0.0877 Japeuthria ferrea 0.501 (12) 0.0982 more surface area for adhesion and more food. Spatial and food resources are expected to be more limited on small hosts, causing intraspecific competition between H. conicus individuals on the same host. Density-depen- dence in growth or survival has been reported in many other organisms (e.g., Begon et al., 1990). Also, the larger hosts may experience less predation pressure, providing a more stable environment. Overall, larger hosts may have higher quality. In contrast, larger H. conicus loads on larger hosts might be due to differences in exposure time to H. con- icus among size-classes of hosts, that is, since larger hosts may be merely older than smaller ones, they might have been adhered to by more and larger (well grown) H. con- icus. To reject this hypothesis, we have to examine Table 5 Correlation coefficients (r) between host size and size of Hipponix conicus. Fisher F to z test: * and ** indicate that the correlation was statistically significant at 5% and 1% level, respectively. Host species r (n) P Thais bronni 0.278 (40) 0.0819 Thais luteostoma 0.041 (247) 0.5180 Omphalius pfeifferi 0.234 (175) 0.0017** Omphalius nigerrima 0.160 (19) 0.5191 Omphalius rusticus —0.024 (255) 0.7042 Chlorostoma argyrostoma 0.177 (52) 0.2109 Lunella coronata 0.226 (80) 0.0434* Astralium haemafragum —0.254 (18) 0.3143 Ergalatax contractus 0.086 (309) 0.1333 Turbo stenogyrum —0.209 (11) 0.5493 Pyrene scripta 0.070 (23) 0.7534 Japeuthria ferrea a — Page 76 The Veliger, Vol. 43, No. 1 Table 6 Mean size (+SD) of snail individuals with at least one Hipponix conicus attached and those without any Hipponix conicus. Mann-Whitney U test: ** denotes the significant difference at 1% level. Mean + SD mm (n) Host species Attached Thais bronni 27.43 + 10.11 (6) Thais luteostoma 24.33 + 6.05 (39) Omphalius pfeifferi 21.44 + 2.96 (36) Omphalius nigerrima 18.44 + 1.27 (9) Omphalius rusticus 20.29 + 1.85 (89) Chlorostoma argyrostoma 2ARO3 se23)7(B1) Lunella coronata 19.99 + 1.03 (48) Astralium haemafragum 20.24 + 2.33 (11) Ergalatax contractus 19.16 + 2.71 (153) Turbo stenogyrum 28.23 + 4.28 (2) Pyrene scripta 15.06 + 0.72 (18) Japeuthria ferrea 30.30 (1) Not attached P 24.38 + 8.35 (3) 0.9028 18.60 + 4.59 (4) 0.241 17.30 (1) 0.3826 14.89 + 4.57 (92) 0.0001 ** 20.62 + 3.22 (36) 0.1102 19.54 + 1.39 (84) 0.0692 17.98 + 4.15 (38) 0.2083 16.74 + 4.32 (214) 0.0001 ** 16.25 + 2.10 (5) 0.0528 14.19 + 1.71 (164) 0.005** 21.96 + 4.36 (11) 0.1924 whether larger hosts are preferred by larger H. conicus disproportionately to their ages. Assuming that the size of each host snail is proportional to its age, it was found that in some host species larger individuals suffered ad- hesion by a significantly larger number of larger H. con- icus than predicted (Yamahira, unpublished data), sug- gesting that differences in exposure time would not be the cause of the differential loads among size-classes of hosts. Matsunaga (1964) stated that H. conicus sometimes moves among host individuals in pursuit of better habi- tats. The extremely biased loads of H. conicus among the species or size-classes of hosts in our study suggests that they actively migrate among host individuals. Although no H. conicus was found living attached on the surface of boulders in the present study, there are also reports of Hipponix being independent of gastropod shells and set- tling on rocks (Matsunaga, 1964, for H. conicus; Mac- pherson & Gabriel, 1962, for H. australis). But, the mi- gration might occur very infrequently because some au- thors have argued that Hipponix is unable to move after settling (Knudsen, 1991; and references therein). In ad- dition, Hipponix might be able to migrate only in the earlier stage of their life cycle because they adapt their shell margin to the configuration of substratum as they grow (Knudsen, 1991). In the slipper limpet, Crepidula adunca, living on the shells of other gastropods and re- producing by direct development like Hipponix, dispersal of newly hatched juveniles has been observed (Putnam, 1964). In conclusion, this study has demonstrated that the dis- tribution of H. conicus was remarkably different among host species or host sizes. These findings suggest that the quality of the snails as a host differs among host species or host sizes. The difference in H. conicus loads could be caused primarily by preferential adhesion to different hosts and/or differential performances on different hosts. Acknowledgments. We would like to thank Dr. W. H. Heard, Dr. K. Nandakumar, Dr. J. Crooks, Mr. N. Takebayashi, and Mr. N. Ota for making useful comments on the manuscript. This is contribution No. 409 from the Amakusa Marine Biological Lab- oratory, Kyushu University. LITERATURE CITED Amio, M. 1963. A comparative embryology of marine gastro- pods, with ecological considerations. Journal of Shimono- seki University of Fisheries 12:15—144. [in Japanese with English summary]. BEGON, M., J. L. HARPER & C. R. TOWNSEND. 1990. Ecology: Individuals, Populations, and Communities. 2nd ed. Black- well: Oxford. CERNOHORSKY, W. O. 1968. Observations on Hipponix conicus (Schumacher, 1817). The Veliger 10:275—280. HaseE, T. 1953. Studies on the eggs and larvae of the Japanese gastropods. Publications of the Seto Marine Biological Lab- oratory 3:161-167. KNUDSEN, J. 1991. Observations on Hipponix australis (Lamarck, 1819) (Mollusca, Gastropoda, Prosobranchia) from the Al- bany area, Western Australia. Pp. 641—660 in F. E. Wells, D. I. Walker, H. Kirkman & R. LETHBRIDGE (eds.), Proceed- ings of the Third International Marine Biological Workshop. The Marine Flora and Fauna of Albany, Western Australia. Vol. II. Western Australian Museum: Perth. Laws, H. M. 1971. Reproductive biology and shell site prefer- ence in Hipponix conicus (Schumacher). The Veliger 13: 115-121. MACPHERSON, J. H. & C. J. GABRIEL. 1962. Marine Molluscs of Victoria. Melbourne University Press: Melbourne. MAatsunaGA, N. 1964. Remarks on the ecological observation of a Japanese hoof shell, Hipponix (Sabia) conicus (Schumach- er). Venus (Japanese Journal of Malacology) 23:149—157 [in Japanese with English summary]. K. Yamahira & F. Yano, 2000 PuTNaAM, D. A. 1964. The dispersal of young of the commensal gastropod Crepidula adunca from its host, Tegula funebral- is. The Veliger 6(Suppl.):63—66. TakabA, Y. 1996. Vertical migration during the life history of the intertidal gastropod Monodonta labio on a boulder shore. Marine Ecology Progress Series 130:117—123. Takapba, Y. & T. Kikucui. 1990. Mobile molluscan communi- ties in boulder shores and the comparison with other inter- tidal habitats in Amakusa. Publications from the Amakusa Page 77 Marine Biological Laboratory, Kyushu University 10:145— 168. YOnGE, C. M. 1953. Observations on Hipponix antiquatus (Lin- naeus). Proceedings of the California Academy of Sciences, Fourth Series, 28:1—24. YonGe, C. M. 1960. Further observations on Hipponix antiquatus with notes on North Pacific pulmonate limpet. Proceedings of the California Academy of Sciences, Fourth Series, 31: 111-119. The Veliger 43(1):78-81 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 Growth and Fecundity of Lymnaea elodes (Gastropoda: Lymnaeidae) under Laboratory Conditions LAUREN FLORIN, BERNARD FRIED! ann ADITYA REDDY Department of Biology, Lafayette College, Easton, Pennsylvania 18042, USA Abstract. Growth and fecundity of Lymnaea elodes were studied under controlled laboratory conditions. Snails maintained in groups of five showed less significant growth (based on shell length) than snails maintained singly at 5 to 29 weeks after the cultures were initiated. This study used newly hatched juveniles that were maintained in artificial spring water at 22°C and fed leaf lettuce ad libitum. Grouped snails fed either leaf lettuce, Tetramin fish food, or freshly killed Helisoma trivolvis snails (FKS) showed maximal growth based on both shell length and body weight only when maintained on lettuce. Submaximal growth was attained on the FKS diet. Snails maintained on Tetramin grew better than those on FKS, but not as well as those on lettuce. Fecundity studies on mature snails maintained either singly or in groups of five on lettuce showed that egg laying was markedly greater at most data points from snails maintained singly. During the course of this study, only four of about 150 snails died in culture suggesting that this lymnaeid is hardy and well suited for laboratory studies. INTRODUCTION Sorensen et al. (1997) identified Lymnaea elodes (Say, 1821) (L. elodes = L. palustris = Stagnicola palustris = S. elodes) as a vector of the ubiquitous trematode Echi- nostoma revolutum (Froelich, 1802) in the USA. Most studies on growth and fecundity of L. elodes are based on observations from natural populations. For example, see the studies of Eisenberg (1966) and Brown et al. (1985). Surprisingly little information is available on growth and fecundity of this snail under controlled lab- oratory conditions. Because this snail is easy to maintain in the laboratory, achieves a shell length of 2 to 3 cm, and is a vector of E. revolutum, it is a useful model for laboratory work. The purpose of this study was to examine the growth and fecundity of L. elodes under controlled laboratory conditions. Specifically, it examined growth and fecun- dity of snails maintained singly versus in groups. It also examined the effects of different diets on snail growth. MATERIALS AND METHODS Lymnaea elodes snails collected from wetlands in north- ern Indiana, USA (Sorensen et al., 1997) provided the original stocks for this study. Snails derived from these stocks were maintained at 22°C in aerated cultures con- taining artificial spring water (ASW) prepared as de- scribed by Ulmer (1970). Each stock tank contained ap- proximately 50 to 100 snails per 5 L of ASW. Snails were ' Author to whom correspondence should be addressed, Phone: 610-330-5463; Fax: 610-330-5705; e-mail: friedb @ lafayette edu maintained under diffuse overhead fluorescent light for 12 hr per day and were fed boiled leaf lettuce (Lactuca sativa) ad libitum. Chalk was added to the cultures as a supplemental source of calcium. Experiment 1 was done to determine the mean shell length of singly raised snails versus those maintained in a group. A single experiment was done using juvenile snails approximately 1 mm in shell length and was set up with five individual or five grouped snails. To avoid pos- sible genetic factors affecting growth, juveniles were se- lected from different egg clutches. Single and grouped snails were maintained in 11-cm diameter finger bowls each containing 150 mL of ASW. Snails were fed leaf lettuce ad libitum, and water and food were changed at least twice a week. Snails were measured with a milli- meter rule, under a dissecting scope, to the nearest 0.2 mm every week for a total of 29 weeks. During the course of these experiments, in which we examined more than 150 snails ranging from | to almost 30 mm in shell length during a span of more than 200 days, mortality was remarkably low in that we had only four deaths. Experiment 2 determined the effects of various diets on snail shell length and wet weight representing five cultured snails per 11-cm diameter bowl in 150 mL of ASW. This study was initiated using mature snails with a shell length of about 14-mm and a wet weight of about 150 mg. These snails were raised in bowls as described above, and fed ad libitum on either leaf lettuce, Tetramin fish food, or freshly killed Helisoma trivolvis snails from which the shell was removed and the body severed in half. The mean shell length of L. elodes was measured every 5 days for 70 days. Additionally, wet weights of L. Florin et al., 2000 Page 79 these snails were also determined at 5-day intervals for 70 days. Experiment 3 was done to determine snail fecundity in isolated versus grouped snails, and used sexually mature snails of about 14 mm in shell length. These snails were maintained in bowls as described for Experiments 1 and 2. The total number of eggs was counted, and eggs were removed from the bowls every 5 days up to day 60. Whenever applicable, the Student’s t-test was used to compare differences in means between populations; and a P value of < 0.05 was considered significant. RESULTS The results of the study (Experiment 1) on growth of single versus grouped L. elodes are presented in Figure 1. Where standard error bars are not shown, they are too small to be included. The shell length increased slowly in both groups until week 4. There was no significant difference (Student’s t-test) in shell length in either group up to week 4. Growth became more rapid in both groups between 5 and 21 weeks. Beyond 21 weeks, growth tend- ed to level off in both groups but more so in single snails. From week 5 to 29, growth of single snails was always significantly greater (Student’s t-test) than that of grouped snails. Growth differences were most apparent at 20 weeks when shells of single snails were 1.5 times longer in shell length than those of grouped snails. The results of the study (Experiment 2) on growth of snails maintained on different diets is shown in Figures 2 and 3. As seen in Figure 2, maximal shell length was noted in snails maintained on leaf lettuce. Snails main- tained on the freshly killed snail (FKS) diet showed the least increase in size. Snails maintained on the Tetramin diet showed growth intermediate between that of the snails maintained on the other diets. By day 18, the shell length of snails maintained on lettuce was significantly greater (Student’s t-test) than that of snails maintained on either Tetramin or FKS. This trend continued for the du- ration of the experiment. Observations from this experi- ment using blotted snail wet weights (weights were de- termined per group of five snails, and averaged to get a weight per snail, precluding the t-test analysis), are shown in Figure 3. Growth here follows the observations seen in Figure 2 in that maximal weight gains were seen in snails fed lettuce, followed by snails fed Tetramin. Least weight gain was seen in snails fed FKS. In fact, in snails fed FKS there was an initial weight loss prior to weight gains beyond day 20. Fecundity data (Experiment 3) are presented in Figure 4. In general, snails maintained singly laid more eggs than those maintained in groups, and only at one data point (day 5) was this trend reversed. Data showing the greater number of eggs per observation period in single versus grouped snails are shown on days 25 and 45 where the number of eggs laid by snails maintained singly ranged from about three to eight times greater than that of grouped snails. Duplicate cultures were used when deaths occurred so that measurements on n = 5 were always possible. Mortalities never occurred in experiments that used single snails. DISCUSSION The low mortality observed herein attests to the hardiness of this snail under controlled laboratory conditions using artificial spring water and frequent changes of food and water. These cultures were non-aerated, and bowls were covered with plastic wrap that had been perforated with needles to facilitate gas exchange. The simplicity of our design will be ideal for future use of this lymnaeid in the laboratory for experimentation. Lymnaeid snails in the laboratory live for approxi- mately 1 year. Data from Hyman (1967) show that lym- naeid longevity varies from 8 to 10 months for L. pal- ustris, and up to 15 months for L. peregra. As reviewed by Hyman (1967), many factors influence the growth of pulmonate snails in laboratory cultures in- cluding snail density, availability of food, and most im- portantly, accumulation of waste products. It is not clear why single L. elodes, beginning at about 5 weeks after culture, showed such growth advantages over those main- tained as a group of five. The amount of water used and the ad libitum feeding would have supplied more than an adequate water volume and food for this species of snail maintained at five per culture. Perhaps substances re- leased into the water by snails maintained five per group had a growth inhibitory effect on snails of this species. Release of excretory-secretory products by pulmonate snails into water (known as snail-conditioned water or SCW) has multiple effects on conspecific snails, includ- ing an influence on intraspecific pairing behavior and at- traction of larval trematodes (Marcopoulus & Fried, 1993). Late in the experiment, the grouped snails seemed to be catching up to the single snails in growth. Reasons for this phenomenon are not known. The effect of diet on lymnaeids has not been thor- oughly investigated, although Pennak (1989) noted that some species of Lymnaea are omnivorous scavengers, whereas others feed on live animals. In our study, L. elo- des fed on lettuce, which is suggestive of a herbivorous diet, and fed on Tetramin, which is suggestive of an om- nivorous diet. L. elodes also fed on freshly killed Heli- soma snails, which is suggestive of a carnivorous diet. Because the growth of L. elodes was optimal on lettuce, the herbivorous diet seems best suited for this lymnaeid, at least under laboratory conditions. Our observations on fecundity in single versus grouped L. elodes did not allow for speculation on the role of self- versus cross-fertilization in this species because snails used in our study came from a sexually mature breeding stock. Therefore, it was not possible to conclude that eggs Page 80 The Veliger, Vol. 43, No. 1 = (= = = £ £& wu Lu ” ” +H +H = <= ~ ~ Oo i=) ce c 2 2 ro o <= <= 2) Y Cc c O 1s) oO o = = 10 15 20 Age in weeks Age in days S$ = Snail diet or FKS diet SS = Single snail T = Tetramin diet GS = Grouped snail L = Lettuce diet 8 8 200 Mean wet weight in mg 8 Mean number of eggs per dish = as TEI Ts a 40 60 Xe 8 ¢ 8B &B R B Age in days Number of days L. Florin et al., 2000 laid by our isolated snails resulted from self-fertilization. However, our findings on enhanced egg production in snails maintained singly in L. elodes support the obser- vations of van Durvenboden et al. (1985) that L. stagnalis maintained singly has a higher oviposition rate than those maintained in groups. LITERATURE CITED Brown, K. M., D. R. DE Vries & B. K. LEATHERS. 1985. Causes of life history variation in the freshwater snail Lymnaea elo- des. Malacologia 26:191—200. EISENBERG, R. M. 1966. The regulation of density in a natural population of the pond snail, Lymnaea elodes. Ecology 47: 889-906. Hyman, L. H. 1967. The Invertebrates, Volume VI: Mollusca I. McGraw-Hill: New York. 792 pp. Page 81 MARCOPOULOS, A. & B. FRIED. 1993. Chemoattraction of Biom- phalaria glabrata (Gastropoda: Planorbidae) for lipid stan- dards and lipophilic factors in leaf lettuce and Tetramin. The Veliger 38:73-74. PENNAK, R. W. 1989. Fresh-Water Invertebrates of the United States: Protozoa to Mollusca. 3rd ed. Wiley: New York. 628 PP. SORENSEN, R. E., I. KANEv, B. FRIED & D. J. MINCHELLA. 1997. The occurrence and identification of Echinostoma revolutum from North American Lymnaea elodes snails. Journal of Par- asitolology 83:169-170. Umer, M. J. 1970. Notes on rearing snails in the laboratory. Pp. 143-144 in A. J. MacInnis & M. Voge (eds.), Experiments and Techniques in Parasitology. W. H. Freeman: San Francisco. VAN DUIVENBODEN, Y. A., A. W. PIENEMAN & A. TER Matt. 1985. Multiple mating suppresses fecundity in the hermaphrodite freshwater snail Lymnaea stagnalis: a laboratory study. An- imal Behavior 33:1184—-1191. Explanation of Figures 1—4 Figure 1. Growth of single (SS) versus grouped (GS) snails. Figure 2. Growth (increase in mean shell length) of snails maintained on various diets. Figure 3. Growth of snails maintained on various diets as described in Figure 2. Growth was based on mean wet weight per snail. Figure 4. Fecundity of single versus grouped snails. THE VELIGER © CMS, Inc., 2000 The Veliger 43(1):82—97 (January 3, 2000) A New Subspecies of the Schoolmaster Gonate Squid, Berryteuthis magister (Cephalopoda: Gonatidae), from the Japan Sea OLEG N. KATUGIN Pacific Research Fisheries Centre (TINRO-Centre), Vladivostok 690600, Russia Abstract. Morphological and genetic variation of the gonatid squid Berryteuthis magister from the North Pacific Ocean is analyzed. Geographical differences were observed between sexually mature squid from different parts of the species range. B. magister from the Japan Sea was clearly distinguished from other conspecific populations by distribution of several morphologic features: dorsal mantle length, nidamental gland length, fin length, and fin width. Genetic di- vergence D, between squid from the Japan Sea and from waters off the Kurile Islands was 0.044, as revealed from 26 protein-coding genetic loci. Significant allele frequency differences were observed at seven of 12 polymorphic loci, and gene differentiation F(ST) equalled 0.12. Data from morphology, genetics, geographical distribution, reproduction, and ecology suggest that B. magister from the Japan Sea constitute a separate taxon of subspecific rank. Berryteuthis magister shevtsovi subsp. nov. from the Japan Sea is described. The subspecies is small, with a large fin, a bicuspid lateral tooth on radula, and weak differentiation between the central and marginal suckers on the tentacular club. It breeds in spring, and produces a small number of large eggs. It leads a bathypelagic life in the Japan Sea at very low temperatures. INTRODUCTION The schoolmaster gonate squid Berryteuthis magister (Berry, 1913) is the type species of the genus Berryteuthis Naef, 1921. Specimens of this squid were first described by Berry (1912) from Puget Sound, and were identified at that time as Gonatus fabricii (Lichtenstein, 1818). Ber- ry noted that they differed from true G. fabricii in several features, the hookless tentacular club being the most no- table. A year later (Berry, 1913), the specimens received specific taxonomic status as Gonatus magister Berry, 1913. Based primarily on tentacular club armature and radula morphology, Naef (1921) placed this gonatid spe- cies in a new genus Berryteuthis. It was not until the discovery and description of several other gonatid species that taxonomic validity of the genus became obvious, and was accepted by malacologists (Okutani, 1968; Roper et al., 1969). Pearcy & Voss (1963) described the species Gonatus anonychus, which shared with B. magister such generic characters as a hookless club and a radula with seven teeth in a transverse row (septemdentate). Okutani (1968) later placed G. anonychus into the genus Berry- teuthis. Berryteuthis magister is presently considered a poly- typic species with two subspecies: B. (m.) magister (Ber- ry, 1913), and the recently described B. (m.) nipponensis Okutani & Kubodera, 1987 (Okutani et al., 1987). The Japanese subspecies is extremely rare, with only three specimens known so far, which were caught in Japanese waters in the Sanriku region (holotype specimen), Toya- ma Bay (Okutani et al., 1987), and off Cape Erimo (Ku- bodera, 1993). All specimens of B. (m.) nipponensis are immature, and can be distinguished from the typical B. magister by the presence of a muscular mantle with small mantle opening (pallial aperture), small fins, and less pro- nounced size differences between central and marginal suckers on the club (Okutani et al., 1987). The nomino- typical subspecies is very abundant in the North Pacific, and has a wide geographical distribution, including mar- ginal basins, such as Japan, Okhotsk, and Bering seas, and the Gulf of Alaska (Nesis, 1987). B. (m.) magister from the Japan Sea is apparently a geographical isolate, which lives under rather specific hydrological conditions (Okiyama, 1993), and exhibits peculiarities both in repro- ductive cycle (Nazumi et al., 1979; Yuuki & Kitazawa, 1986; Natsukari et al., 1993), and morphology. The analysis of certain morphologic characters, togeth- er with the analysis of inherited molecular variation as revealed by protein electrophoresis, provided reliable ev- idence in favor of taxonomic separation of B. (m.) mag- ister from the Japan Sea on a subspecific level when com- pared with B. (m.) magister from other parts of its distri- bution range. A new subspecies of B. magister is de- scribed. ; MATERIALS AND METHODS A total of 3501 specimens of squid were used for the analysis of intraspecific differences. All were morpholog- ically identified as B. (m.) magister, hereafter referred to as B. magister (Figure 1, Table 1). Three thousand one hundred and seventeen specimens in 14 sample lots were taken for the size composition comparison of B. magister from different regions of the North Pacific Ocean: Japan O. N. Katugin, 2000 28 _4 NKU9, NKULO Oo ¥g@ Se) Co) CKUS KUN CK | o -@ Es SKU? JAP1, JAP2, JS Page 83 BER14, BS1, BJ Figure | Berryteuthis magister. Location of samples in the North Pacific Ocean. Sea (Kitayamato Bank); Pacific waters off the Kurile Is- lands; the Kommander Islands (central bank); the north- western Bering Sea (Olutorsko-Navarinskyi Region); and the northeastern Pacific (central part of the Aleutians and Gulf of Alaska). All specimens were sexually mature, which suggests that squid were taken from breeding stocks. Squid from the Japan Sea and the northwestern Pacific which were subjected to size composition analysis were caught during 1988, whereas those from the north- eastern Pacific Ocean were caught in 1985 and 1987. One hundred and sixty specimens in seven sample lots were taken for the analysis of fin proportions of immature and mature B. magister from the North Pacific Ocean. Morphology of the radula was studied in mature spec- imens from the Japan Sea (25 specimens), and in mature and immature specimens from the northwestern Bering Sea (25 specimens) under light microscope and under scanning electron microscope (SEM) JEOL JSM-25SII. Soft tissues of the buccal mass were macerated in 10% NaOH for 24 hr. The radular ribbon was then removed, washed in tap water, and placed in 70% ethanol. Radular structures were resolved under SEM with accelerating voltage of 25 kV, and with a magnification of 70x. A total of 234 B. magister specimens in two sample lots, representing two geographical regions, the Japan Sea, and the northwestern Pacific, were taken for electro- phoretic comparison of enzyme-coding genetic loci. De- tails of multilocus electrophoretic analysis for the squid are described elsewhere (Katugin, 1991, 1992, 1993, 1995, in press; Katugin et al., 1992). Abbreviations of genetic loci, revealed by protein electrophoresis, were for the most part conventional, and were used following gen- eral recommendations suggested by Allendorf & Utter (1979). Basic calculations were made using a BIOSYS-1 computer program (Swofford & Selander, 1981, 1989). The following standard variability measures were used: mean sample size per locus, mean number of alleles per locus, two criteria of polymorphism (0.95 and 0.99), and mean heterozygosity per locus (biased, unbiased, and di- rect-count estimates). Contingency comparison was con- ducted by chi-square test (Workman & Niswander, 1970), and genetic differentiation was assessed using F(ST) (Wright, 1978). Standard genetic distance Dy (Nei, 1972) was used to measure proportion of codon differences be- tween geographical populations. Estimation of presump- tive intersubspecific divergence time (t) was made using Nei’s formula: t = 5* 10°D,y (Nei, 1987). Specimens used in the description of the new subspe- cies were fixed in a 5% formaldehyde solution for 7 days followed by transfer to 45% isopropanol solution for long-term preservation. RESULTS Size Composition Mature specimens of B. magister exhibited a geograph- ical pattern with regard to size: dorsal mantle length (DML) range and mean for males and females were the Page 84 The Veliger, Vol. 43, No. 1 Table 1 Berryteuthis magister: Locations, dates, and number of sampled specimens. Abbreviations: AM—mature adults; AI— immature adults; JU—juveniles; *—samples for size composition analysis; **—-samples for fin dimensions analysis; ***__samples for multilocus electrophoretic analysis. Sample Region Coordinates Date Number Maturity Japan Sea JAP1* Kitayamato Bank 39°56'N 133°53’E 8.02.1988 434 AM JAP2* Kitayamato Bank 39°59'N 133°49/E 12.04.1988 137 AM GRRE Russian Primorie 42°30'N 133°42'E 23.05.1996 104 AM, AI North-eastern Pacific Ocean ALA3* Gulf of Alaska 55°27'N 155°46'W 27.06.1985 24 AM ALA4* Gulf of Alaska 55°37'N 155°25'W 26.06.1986 15 AM ALES5* Aleutian Islands 53°15'N 165°20'W 13.06.1985 89 AM ALE6* Aleutian Islands 53°43'N 165°10'W 15.06.1987 ii AM AL** Aleutian Islands 53°05'N 171°36'W 24.03.1989 20 AM, AI North-western Pacific Ocean Kurile Islands SKU7* Southern Kuriles 44°48'N 148°53’E 15.05.1988 168 AM CKU8* Central Kuriles 46°51'N 152°06’E 11.05.1988 29 AM KU** Central Kuriles 46°50'N 152°06’E 14.09.1996 43 Al CK*** Central Kuriles 47°12'N 152°29'E 8.11.1991 133 Al NKU9* Northern Kuriles 49°46'N 154°57'E 19.06.1998 95 AM NKU10* Northern Kuriles 49°40'N 156°62'E 7.07.1988 223 AM OJ** Okhotsk Sea 45°23'N 145°35'E 2.06.1988 17 JU Kommander Islands KOM11* Central Bank 54°30'N 167°05’E 28.01.1988 202 AM KOM12* Central Bank 54°24'N 167°06'E 22.05.1988 407 AM KOM13* Central Bank 54°29'N 167°02'E 1.12.1988 1154 AM Bering Sea BER14* Navarin Region 61°40'N 176°30'E 17.09.1988 129 AM BS1** Navarin Region 61°23'N 175°23'E 4.09.1996 20 AM BJ** Navarin Region 61°43'N 176°36'E 9.09.1996 23 JU BS2** Olutorsky Bay 60°02'N 168°38'E 8.09.1996 20 AM lowest in samples from the Japan Sea (Figure 2). Mean size for males was 150.2 + 0.7 mm in February, and almost the same, 151.5 + 3.8 mm, in April. Females in February were smaller than in April, having a mean DML of 180 + 0.7 and 192.2 + 1.7 mm, respectively. Size differences were significant (P < 0.001), which could be explained by a higher proportion of mated females in April (84%) than in February (11%). Mated females with traces of copulation (ropes of spermatangia implanted in the inner wall of the mantle) had a mean size of 189.7 + 1.8 mm in February and 193.8 + 2.0 mm in April, show- ing no significant differences. In the western and south- western Japan Sea, DML values for copulated females were almost as low as in the central part, with a mean value of 195.6 + 2.5 mm for specimens from the Oki Bank (based on data from Nazumi et al., 1979). In the northeastern and northern Japan Sea off the Russian Pri- morie coast, mature squid were somewhat larger, with DML values of 177.1 + 4.0 mm for males and 207.6 + 3.8 mm for females (P. P. Railko, personal communica- tion). Among 440 prespawning females sampled by Dr. G. A. Shevtsov in February through May 1988 from the Kitayamato Bank, seven (1.6%) specimens had DML larger than 220 mm, one of them with a maximum value of 315 mm. Mean DML for copulated females was 194 mm, with a mode of 202 mm. Among 320 ripe males, eight (2.5%) had DML larger than 170, and the maximum value for one of them was 202 mm. Geographical trends in size distribution were also ob- served for B. magister from the northwestern Pacific Ocean: the largest specimens were present in the southern part of the region, and mean size generally decreased northward (Figure 2). Both males and females caught off the southern Kuriles were very large with mean DML values of 245.9 + 1.8 and 313.7 + 6.3 mm, respectively. The smallest males with mean DML of 205.1 + 1.5 mm were from the northern Kuriles, and the smallest females with mean DML of 253.3 + 1.3 mm were from the Be- ring Sea. Squid from the northeastern Pacific region were characterized by their very large size when compared O. N. Katugin, 2000 Males —]—_- JAPI —1} JAP2 ——_{]}__ SKU? === CKUB == NKUQ — NKU10 —_]-____- KOM1I1 —__- KOMI2 —]}#—____- KOM13 —]—_- BER14 — Bo ALA3 ALA4 — a= ALES ALE6 DML, mm Page 85 Females —_j]— JAPI —_]}—__—_- JAP2 — Sala SKU? ——S aa CKU8 ee NKUQ a NKU10 = KOM11 ———_—_—_ I ————____- KOM12 ——_]___——_- KOM13 —}—_ BERI4 —— f2)}-— ALA3 —{_]_ #—___ a4 ———_{}—__ ALES SS aS ALE6 DML, mm Figure 2 Berryteuthis magister. Size distributions of mature males and females from the North Pacific Ocean. DML, dorsal mantle length; vertical line, mean; rectangle, 95% confidence limits; horizontal line, range. with conspecific populations from the other regions. In the northeastern Pacific Ocean, the largest males were collected off the Aleutian Islands with mean DML of 257.5 + 6.1 mm, and the largest females were found in the Gulf of Alaska with mean DML of 301.1 + 11.2 mm. In a sequence of regions—the Japan Sea, the northwest- ern, and northeastern Pacific-respective mean values of DML were 150.3 + 0.7, 224.9 + 0.6, and 242.9 + 4.5 mm for males, and 184.2 + 0.8, 273.5 + 0.9, and 294.0 + 3.0 mm for females. Size distribution graphs for mature B. magister re- vealed some regional-specific features (Figure 3). One of the most striking features is a gap between size distri- bution curves for specimens from the Japan Sea and from other regions of the North Pacific Ocean. DML distribu- tions for males and females from the Japan Sea are almost unimodal with modal classes 150—160 and 180-190 mm, respectively. Males larger than 190 mm, and females larg- er than 220 mm were extremely rare. In the northwestern Pacific there are main modal classes of 220 mm for males, and 270 mm for females with a pronounced pro- portion of large animals. Total share of females with DML larger than 300 mm constituted up to 20% of all ripened females in the spawning stock off the Komman- der Islands. In the northeastern Pacific unimodality of size distribution is even less evident. Two size groups of B. magister were present in different ratios in the Aleutians and in the Gulf of Alaska: small squid with equal modal classes for males and females of 210 mm, and large squid with modal classes of 270 mm for males and 300 mm for females. On the scatterplot of DML against nidamental gland length (NGL), a geographical pattern is also seen. Mea- surements of both characters separated sexually mature squid from the three regions of the North Pacific Ocean (Figure 4). Squid from the Japan Sea occupy a distinct separate zone on the graph, while distribution patterns for specimens from the northwestern and northeastern Pacific overlap considerably. Mean DML values for B. magister from the Japan Sea, northwestern, and northeastern Pa- cific were, respectively, as follows (values for males are given in parentheses): 184.2 + 0.8 (150.3 + 0.7), 273.5 + 0.9 (224.9 + 0.6), and 294.0 + 3.0 (242.9 + 4.5) mm. Mean NGL values for females were 78.2 + 0.5, 107.1 = 0.4, and 126.2 + 1.7 mm, respectively. Line, or simple regressions, of NGL by DML, for squid from all the three North Pacific regions are as follows: 5.91 + 0.39 * DML (Japan Sea), 45.0 + 0.23 * DML (northwestern Pacific), and 61.7 + 0.22 * DML (north- eastern Pacific). First coefficients do not vary signifi- cantly between the last two regions, and were signifi- cantly different (P < 0.0001) from that of the Japan Sea. Page 86 number 300 - 250 Males 200 - + Sea of Japan 150 - 3 North— western Pacific -e North — eastern Pacific 100 - / / i a | \ / A NG o -|\—1+ of, : ———— 100 400 500 number 140 } 120 /| IN a Females mf 100 f ) at) | +Sea of Japan | 3 North — western Pacific 60 | | p -e- North — eastern Pacific | iy] aa | 20 ) o-@ Ske sk Hi rt See @Mi- = Z o-2 Se 100 200 300 400 500 DML, mm Figure 3 Berryteuthis magister. Size distributions for mature males and females from three regions in the North Pacific Ocean. Second coefficient, which shows the angle of the regres- sion line, is almost 1.5 times higher in Japanese samples than in the others, meaning that NGL is increasing more rapidly with DML in mature squid from the Japan Sea. This could be explained by the fact that the size distri- bution curve in mature B. magister from the Japan Sea is narrow with only one predominant class, whereas in the other regions, size distribution curves are wide, and composed of several size classes. Fin Measurements and Indices Measurements of fin length (FL) and fin width (FW), and indices of FL/DML and FW/DML varied consider- ably among samples of squid in different onthogenetic stages, and from different geographic regions. For the samples from the northwestern Pacific the val- ue of FL/DML rapidly increases in juveniles from 0.401 + 0.006 (sample JB, DML = 54.9 + 1.3) to 0.471 + 0.007 (sample JO, DML = 100.1 + 4.0), reaches a max- imum of 0.495 + 0.003 in immature adults (sample KU, DML = 197.0 + 3.1), and slowly decreases to 0.489 + 0.004 in prespawning squid (samples BSI and BS2, DML = 250.7 + 3.3) (Figure 5). Aleutian adults (sample AL, The Veliger, Vol. 43, No. 1 NGL, mm 190 140 90 4 a anid av ene a AON ow 40 /er + 140 DML, mm NGL, mm B 190 | | 140 | | —= i, | | 90 | SS | 4 — Sea of Japan | O — North— western Pacific ® — North—eastern Pacific 40 + T T T T T 140 190 240 290 340 390 DML, mm Figure 4 Berryteuthis magister. Relationship between dorsal mantle Jength (DML) and nidamental gland length (NGL) in mature females from three regions in the North Pacific Ocean. A. Distribution of individual values. B. Means and ranges by regions. DML = 218.2 + 12.4) have the shortest fins with a mean FL/DML index of 0.478 + 0.004, while adults from the Japan Sea (sample JS, DML = 155.7 + 6.7) are charac- terized by the highest average value of this index, 0.501 + 0.006. Adult specimens from the Japan Sea lie between juveniles and immature adults on the graph of the FL index plotted against DML (Figure 5B). For the samples from the northwestern Pacific the val- ue of FW/DML slightly increases in juveniles from 0.759 + 0.005 (sample JB) to a maximum of 0.805 + 0.008 (sample JO), with a subsequent slow decrease through 0.718 + 0.005 in immature adults (sample KU) to a min- imum of 0.0.693 + 0.004 in prespawning squid (samples BS1 and BS2) (Figure 6). Aleutian adults (sample AL) have a mean index value of 0.748 + 0.008, which is intermediate between juveniles and immature adults from the northwestern Pacific. Adults from the Japan Sea (sam- ple JS) are characterized by a mean FW/DML index of 0.771 + 0.018, which falls between mean values for sam- ples of juveniles and immature adults on the graph of this index plotted against DML (Figure 6B). Radula The radula of B. magister is of the heterodont type, and is characterized by seven teeth in a transverse row (versus five teeth in most other gonatids): rachidian tooth R, two lateral teeth L, two inner marginal teeth M1, and two outer marginal teeth M2. Based on nomenclature for O. N. Katugin, 2000 FL/DML 0.6 | A 0.5 - & o ace ED oe 4 “halle Sie: o ‘i 00 oS O m9 ia) 0.4 Sth, ty 0.3 02 4 — Sea of Japan Ohl = © — North—western Pacific e — North—eastern Pacific ORS, Teen T WY Tage oat T UST u ial eon ae ee ae 0 50 100 150 200 250 300 350 DML, mm FL/DML 0.6 7 B 0.5 | 0.4 + 0.34 QA= | OMS | 4 — Sea of Japan 9 — North—western Pacific bp , 0) 50 100 150 200 250 300 35 DML, mm Figure 5 Berryteuthis magister. Relationship between fin length index (FL/DML) and dorsal mantle length (DML). A. Distribution of individual values. B. Means and ranges by regions. cephalopod radulae, suggested by Nixon (1995), the rad- ular formula for B. magister is represented as: R, -, L, M1, M2. There are certain differences in radular morphology be- tween B. magister from the Japan Sea, and from the northwestern Bering Sea (Figure 7). The rachidian tooth is essentially invariant, and consists of a mesocone with one lateral cusp on either side in all 50 adult animals studied. The lateral tooth on each side of the rachidian has one central cusp and one outer cusp in 22 of 25 spec- imens studied from the Japan Sea. In 18 of 25 studied specimens from the northwestern Bering Sea, there were three cusps on each lateral tooth: one large central cusp, One outer cusp, and a very small inner cusp. Three spec- imens from the Japan Sea had a tricuspid lateral, while seven specimens from the northwestern Pacific had a di- cuspid lateral. None of the Japan Sea squid had a well- developed inner cusp on L. There was no visual corre- lation between a development of this cusp and a maturity stage in squid from any of two regions. All marginal teeth Page 87 FW/DML A i elay n 08 §) oo o, Ae ogy? (Bh, ee a a ee ° 4 Feo oo? 0 ° a & §) Snyact a oe 06 - 0.4 4 Or23— 4 — Sea of Japan Oo — North—western Pacific e — North—eastern Pacific o + poo a ie} 50 100 150 200 250 300 350 DML, mm FW/DML 41 ~ B 0.8 ! =F 0.6 - 0.4 - O25 | 4 —Sea of Japan 9 —North—western Pacific o +— ana pat 0 50 100 150 200 250 300 350 DML, mm Figure 6 Berryteuthis magister. Relationship between fin width index (FW/DML) and dorsal mantle length (DML). A. Distribution of individual values. B. Means and ranges by regions. in all specimens studied were unicuspid, as they are in most other Recent, and even fossil cephalopods (Nixon, 1995). Typical radular formulas for B. magister from the two different geographic regions would be: 5R;, -, Ly, M1,, M2, (for most of the specimens from the Japan Sea), >R;, -, L;, M1,, M2, (for most of the specimens from the northwestern Bering Sea). Genetic Evidence Genetic variation and differentiation between B. mag- ister from the Japan Sea, and from the northwestern Pa- cific Ocean were assessed from the analysis of 26 putative genetic loci, coding for the enzymes and a polymorphic general protein zone. The following 12 loci had more than one allele present in at least one of the two samples: Aat-2 (four alleles), Est-2 (four alleles), Fh-/ (five al- leles), Jdh-1 (four alleles), Ldh-2 (four alleles), Mdh-2 (four alleles), Mpi-/ (three alleles), Pep-2 (four alleles), Ped (five alleles), Pgm-/ (four alleles), Pnp (three al- leles), Ugp (two alleles) (Table 2). Fourteen loci were monomorphic in the Japan Sea sample, and 15 in the central Kurile sample. Genetic variability measures did not vary significantly between sampled populations from the Japan Sea and the Kurile region (Table 3). On aver- Page 88 The Veliger, Vol. 43, No. 1 Table 2 Berryteuthis magister: Genetic loci, alleles and allele frequencies for squid from the Japan Sea (sample JS) and north-western Pacific Ocean (sample CK). Number of specimens screened for each locus are given in brackets. Locus Sete Locus Se Allele JS CK Allele JS CK Aat-2 (51) (130) Idh-1 (102) (130) 125 0.020 0.008 160 0.064 0.038 112 0.098 0.112 120 0.505 0.262 100 0.873 0.881 100 0.387 0.654 88 0.010 0.000 85 0.044 0.046 Agp (104) (130) Idh-2 (104) (130) 100 1.000 1.000 100 1.000 1.000 Ck (104) (20) Lap-1 (104) (20) 100 1.000 1.000 100 1.000 1.000 Est-1 (104) (130) Lap-2 (104) (20) 100 1.000 1.000 100 1.000 1.000 Est-2 (104) (127) Ldh-2 (103) (130) 120 0.000 0.024 130 0.024 0.008 100 0.063 0.972 100 0.398 0.550 92 0.933 0.004 70 0.034 0.000 80 0.005 0.000 40 0.544 0.442 Fh-1 (99) (130) Mdh-1 (104) (130) 155 0.061 0.046 100 1.000 1.000 125 0.086 0.046 Mdh-2 103 107 100 0.798 0.873 ; ue) ( i 90 0.056 0.031 115 0.073 0. a 40 0.000 0.004 100 0.587 0.528 75 0.340 0.341 Gap (104) (20) 58 0.000 0.009 100 1.000 1.000 age, almost 40% of loci were polymorphic, and the ob- served heterozygosity values ranged from 0.128 in the Japan Sea to 0.137 in the Kuriles, which corresponded to the estimates, revealed earlier on a different sample of Locus Se SE Locus See Allele JS CK Allele JS CK Me (104) (20) Pgm-1 (63) (133) 100 1.000 1.000 108 0.056 0.008 105 0.175 0.120 Mpi-1 104 130 Pe a) 30) 100 0.746 0.805 120 0.034 0.000 97 0.024 0.068 100 0.962 1.000 75 0.005 0.000 Pk-1 (104) (20) Mpi-2 (104) (130) 100 1.000 1.000 100 1.000 1.000 Pk-2 (104) (20) Reo (104) (130) 100 1.000 1.000 100 1.000 1.000 Pnp (103) (130) 120 0.141 0.300 Pep-2 101 125 a z Wine z ae 100 0.845 0.688 : 51 0.015 0.012 120 0.010 0.028 100 0.901 0.904 Sod (104) (130) 20 0.035 0.068 100 1.000 1.000 Pgd (103) (133) Ugp (111) (130) 103 0.019 0.026 100 0.847 0.781 100 0.961 0.932 85 0.153 0.219 98 0.010 0.030 95 0.010 0.004 92 0.000 0.008 loci in B. magister from the northwestern Pacific (Katugin, 1993). Significant heterogeneity between the Japan Sea and Kurile samples of B. magister was revealed at seven of 12 loci: Est-2, Idh-1, Ldh-2, Mpi-1, Pep-2, Pgm-1, and Table 3 Berryteuthis magister: Variability measures at 26 genetic loci in sampled populations of squid from the Japan Sea (sample JS) and north-western Pacific Ocean (sample Ck). Population samples Variability measure JS CK Mean sample size per locus 100.1 + 2.5 99.4 + 9.7 Mean number of alleles per locus 22) 1053 Poel a (0,3) Percentage of polymorphic loci (0.95 criterion) 38.5 38.5 (0.99 criterion) 46.2 42.3 Mean heterozygosity per locus (biased estimate) 0.140 + 0.038 0.135 + 0.038 (unbiased estimate) 0.141 + 0.039 0.135 + 0.038 (direct-count estimate) 0.128 + 0.035 0.137 + 0.039 O. N. Katugin, 2000 Table 4 Berryteuthis magister: Contingency comparison and fix- ation index F(ST) of squid from the Japan Sea and north- western Pacific Ocean. Abbreviations: df—degrees of freedom; Na/—number of alleles; P—probability for the chi-square test; *—-significant F(ST) value. Locus Nal Chi-square df P F(ST) Aat-2 4 3.620 3. 0.30554 0.001 Est-2 4 408.086 3. 0.00000 0.825% Fh-1 5 6.427 4 0.16947 0.007* Idh-1 4 34.992 3 0,00000 0.057% Ldh-2 4 18.859 3. 0.00029 0.016% Mdh-2 4 5.137 3. 0.16203 0.003 Mpi-1 3 10.174 2 0.00618 0.017% Pep-2 4 17.793 3. 0.00049 0.006 Ped 5 4.807 4 0.30763 0.003 Pgm-1 4 13.866 3 0.00309 0.007% Pnp 3 16.474 2 0.00026 0.034* Usp 2 3.414 1 0.06465 0.007 Totals 543.648 34 0.00000 0.120% Pnp (Table 4). Significant F(ST) values were observed at the following seven loci: Est-2, Fh-1, Idh-1, Ldh-2, Mpi- 1, Pgm-1, and Pnp—and mean total value for 12 poly- morphic loci was 0.12 (Table 4). Genetic divergence Dy between the two sampled populations of B. magister amounted to 0.044. DISCUSSION Comparative data from morphology (size distribution, fin measurements, radula), and genetics (electrophoretically detectable protein-coding loci) in B. magister revealed a pattern of intraspecific variation, which could be ex- plained in terms of taxonomic and evolutionary relation- ships. Two independent sources of information showed that the population(s) from the Japan Sea differs signifi- cantly from conspecific populations in other regions of the North Pacific Ocean. Among morphologic indicators of the Japan Sea populations are: small body dimensions as revealed by distribution of DML, unimodality of size distribution for either sex, a comparatively large fin, and a radula with bicuspid lateral teeth. The small size of B. magister from the Japan Sea is in good agreement with growth data as revealed by statolith microstructure analysis of specimens from the Rebun Bank (Japan Sea), and from the Kitamyamato Bank (Okhotsk Sea). Squid from the Japan Sea are character- ized by a slow growth rate (based on growth curves), and by comparatively large hatchlings (based on two main radii of the natal ring) (Natsukari et al., 1993). Ripe oo- cytes from oviducts of mature females from the Japan Sea had a maximum diameter of 5.94 mm (Nazumi et al., 1979), which is almost 1.5 times larger than the size of oocytes (4.2 mm) from the western Bering Sea specimens Page 89 (Fedorets & Kozlova, 1986). Small body size and large oocytes result in low individual fecundity of B. magister from the Japan Sea, with a maximum number of eggs of nearly 4000 per female (Nazumi et al., 1979), while the maximum number of eggs per ripe female in the western Bering Sea amounted to 29,000 (Fedorets & Kozlova, 1986), which is almost seven times higher. Comparatively low individual fecundity of the species in the Japan Sea could be due to onthogenetically involved reproductive features, such as dynamics of oocyte formation and de- velopment. It was shown that the germinative epithelium with gonial cells continues to produce future oocytes in fully mature females of B. magister from the northwest- ern Pacific (Reznik, 1982). Small size of females from the Japan Sea could be due to a slow growth rate and/or early maturation. If the latter is the case, then egg pro- duction and consequently, individual fecundity will be lower in smaller females from the Japan Sea, and higher in larger females from the northwestern Pacific. Low fe- cundity, combined with the production of eggs with a higher yolk content, (the increase of egg diameter from 4.2 to 5.94 mm theoretically results in a nearly three-fold increase of yolk content) should lead to an increased sur- vival rate for paralarvae in the rather specific oceanologic conditions of the Japan Sea. Higher survival rates for an- imals with a higher content of egg yolk is due to the so- called effect of embryonization (Zakhvatkin, 1975). In such cases, a hatchling is born with more developed fea- tures, and hence more prepared for life, than a hatchling with lower yolk content. Reproductive strategies of B. magister from the Japan Sea encompass, particularly, time and longevity of spawning season. In this region, the spawning season of squid is more restricted in time, judging by the period of hatch, assessed from statoliths analysis (Natsukari et al., 1993), and is confined to spring with a single peak (Yuuki & Kitazawa, 1986). In the northwestern Pacific, spawning of B. magister is much more extended, and covers a long period from summer through autumn, terminating in winter, with at least two peaks (Fedorets, 1983; Kubodera, 1982, 1992; Nesis, 1995). Squid from the northwestern and northeastern Pacific are hardly distinguishable based on external morphology (radulae have not been studied in specimens from the northeastern Pacific). Distributions of DML and NGL could be measured by a Coefficient of Difference, CD,— a formal descriptor of a subspecific differentiation be- tween conspecific populations (Mayr, 1969). The critical value of subspecific differentiation is at CD of approxi- mately 1.28, when 90% of specimens from one popula- tion differ from 90% of specimens from the other popu- lation. CD values between specimens from the north- western and northeastern Pacific were 0.37 for DML, and 0.66 for NGL, which is considerably lower than the crit- ical value of 1.28, and corresponds to the maximum non- overlap of 74% between character distribution curves. At Page 90 The Veliger, Vol. 43, No. 1 Table 5 Intraspecific normalized genetic identity, I, and standard genetic distance, Dy (Nei, 1972), for different species of squid (L—number of genetic loci). Evolutionary Taxon divergence L Illex argentinus populations 25 sibling species 25 Martialia hyadesi populations 39 sibling species 39 Todarodes pacificus populations 11 Loligo vulgaris subspecies 18 Loligo forbesi populations 33 geographical populations 33 Loligo gahi populations 21 Berryteuthis magister populations 4 populations 19 geographical populations 4 (subspecies) 26 the same time, specimens from the Japan Sea differ from the others with CD values of 2.28 for DML, and 1.47 for NGL, which corresponds to more than 93% of nonoverlap between character distribution curves. Gaps in distribu- tions of each of the two morphometric characters suggest that B. magister populations from the Japan Sea differ from conspecific populations from other parts of the spe- cies range on a subspecific level. It was shown earlier that genetic differences between spatially isolated populations of B. magister from the northwestern Pacific, as revealed by Nei’s (1972) genetic distance (Dy), were very small, of 0.001 (Katugin, 1993). Genetic divergence between conspecific populations of B. magister from the Japan Sea and from Pacific waters off the Kuriles appeared to be 44 times higher: Dy = 0.044. This estimate is slightly higher than the genetic distance between two African subspecies of Loligo: L. (vulgaris) vulgaris, and L. (v.) reynaudii (Augustyn & Grant, 1988) (Table 5). When approximated on a time scale, the esti- mate of Dy, = 0.044 corresponds to the time of evolu- tionary divergence between populations of 220,000 years (Nei, 1987). It means that B. magister populations in the Japan Sea were isolated from the ancestral northwestern Pacific conspecific populations during the Riss glaciation period (Meso-Pleistocene, 250,000—125,000 years ago). Presumably, during the Riss and Vurm Periods, the Japan Sea Basin was disconnected from adjacent northwestern Pacific waters. During one of the latest interglacial trans- gressions of the sea level, the Japan Sea territory was colonized by various species of the boreal Pacific fauna through a channel through the middle of modern Sakhalin Island (Nishimura, 1964). Colonization events probably included ancestors of B. magister. Chains of islands, which were formed during sea level regressions, likely served as barriers to gene flow between founder and pa- rental populations. Apparently, these barriers were effec- I Dy Reference 0.979—1.000 0.021—0.000 Carvalho et al. (1992) 0.878—0.904 0.130-0.101 Carvalho et al. (1992) 0.998-0.999 0.002-—0.001 Brierley et al. (1993) 0.508—0.509 1.790-1.775 Brierley et al. (1993) 0.997—1.000 0.003—0.000 Kim (1993) 0.970 0.030 Augustyn & Grant (1988) 0.998 0.002 Brierly et al. (1993) 0.898—0.902 0.108-0.103 Brierly et al. (1993) 0.997—1.000 0.003—0.000 Carvalho & Pitcher (1989) 0.989—1.000 0.01 1—0.000 Katugin (1995) 0.999 0.001 Katugin (1993) 0.980—0.998 0.020—0.002 Katugin (1995) 0.957 0.044 (this paper) tive for a nectobenthic bathypelagic species such as B. magister. Hydrological conditions in the recently formed sea basin, which were different from those of the north- western Pacific, served as strong selective forces. As a result, a specific ecomorphotype of B. magister evolved with an effective reproductive strategy. Low temperature and geographic isolation of squid in the Japan Sea were presumably among possible triggers for maturation at smaller size, and consequently for evolving of a pedo- morphic line of B. magister. Pedomorphic subspecies are known among living and fossil animals, and are most likely to be formed on a periphery of a species range as a response to regular changes in environment (Eldredge & Gould, 1972; Raff & Kaufman, 1983). Interpopulation (intersubspecific) genetic differences were acquired dur- ing a long and effective geographic isolation of B. mag- ister in the Japan Sea. The absence of gene flow between the Japan Sea and adjacent B. magister populations is confirmed by the high F(ST) value of 0.12, and corre- sponding estimate of theoretical number of only two mi- grants per generation. It means that two specimens per generation are required to maintain the observed value of genetic differentiation. For very large effective popula- tion sizes of millions of animals, the estimate of two mi- grants in each generation means that there is almost no contact (gene exchange) between populations. At the same time, a Dy value of 0.044 is comparatively small and does not indicate that these populations represent two different species. Typically, for sexually reproducing or- ganisms with an allopatric model of speciation, which is most likely to be the case in B. magister, Nei’s (1972) genetic distance values between closely related species fall around 0.7—0.8 (Ayala, 1983). All facts suggest that populations of B. magister from the Japan Sea have acquired a subspecific level of evo- lutionary and taxonomic divergence. Subspecific status of O. N. Katugin, 2000 M2; is} Mi, om L(2) iRy \ Figure 7 Radula of Berryteuthis magister. A. Maturating male (stage II; 214 mm DML) from the western Bering Sea, SEM photo, mag- nification 70X. B. Diagram for the same specimen. C. Mature male (stage V, 134 mm DML) from the Japan Sea, diagram. B. magister from the Japan Sea satisfies the following criteria: (1) geographical isolation; (2) specific habitat (hydrological conditions); (3) characteristic traits in gen- eral morphology (body size, fin proportions, club arma- ture, radula); (4) certain peculiarities in reproductive traits (egg size and fecundity) and behavior (spawning time and dynamics); and (5) significant differentiation at structural genetic loci. DESCRIPTION or A NEW SUBSPECIES oF B. MAGISTER Based on morphological, genetic, reproductive, and eco- logical data, I propose a new taxon of a subspecific rank. The formal description for a new subspecies of the gon- atid squid B. magister from the Japan Sea is presented below. Page 91 Table 6 Berryteuthis magister shevtsovi Katugin, subsp. nov: Principal measurements (mm) and indices for holotype and paratype. Diagrammatic illustrations of measure- ments are given in Figures 8, 10, and 11. Character Holotype Paratype Catalog No BMJSOO1 BMJS002 sex male female maturity stage Vv Ill DML 162 172 MW (MWI) 59 (36.4) S71 B31) FL (FLI) 79 (48.8) 86 (50.0) FW (FWI) 134 (82.7) 138 (80.2) HW (HWI) 41 (25.3) 33 (19.2) ED (EDI) 22 (13.6) 20 (11.6) TTL (TTLI) 255 (157.4) 231 (134.3) SZL 117 _- CLDL 71 — CLVL 51 — CLWL 21 19 CLMW 11 9 AI AL (AILD 99 (61.1) 94 (54.7) A 8 8 B 14 16 Cc 49 48 D 28 22 All AL (ATILI) 107 (66.0) 110 (64.0) A 15 14 B 16 16 C 53 50 D 23 30 Alll AL (AINLD 114 (70.4) 108 (62.8) A Lz 16 B 12 13 Cc 59 54 D 26 25 AIV AL (AIVLD) 105 (64.8) 99 (57.6) A 15 14 B 16 — HCL 31 — D 28 — AIV FA 105 — Berryteuthis magister shevtsovi Katugin, subsp. nov. (Figures 7-11, Tables 6, 7) Material examined: Seven adult specimens were screened for variation in morphologic traits; two of them were in fairly good condition, and hence selected as type specimens. Holotype: male (mature, stage V), 162 mm DML; paratype: female (maturating, stage I), 172 mm DML (from the same haul as holotype). Deposition of the type specimens: collection of cephalopods, laboratory for squid research, Pacific Research Fisheries Centre (TIN- Page 92 The Veliger, Vol. 43, No. 1 Table 7 Comparison of character states for different subspecies of Berryteuthis magister. Abbreviations: DML—dorsal mantle length; NGL—nidamental gland length; FLI—fin length index; FWI—fin width index; radula teeth: R—rachidian, L— lateral, M1—first marginal, M2—second marginal; d(centr)—diameter of largest sucker ring (central region of the ma- nus); d(marg)—diameter of smallest sucker ring (marginal region of the manus). Character Maturity (stage) DML—males (mm) mean DML—females (mm) mean NGL (mm) mean FLI mean FWI mean Differences between central and marginal club suckers d(centr)/d(marg) Typical radula lateral tooth Maximum individual fecundity (number of eggs) Largest egg diameter (mm) Sea temperature (°C) Ecotype Breeding season shevtsovi (new) mature (IV—V) Subspecies magister nipponensis (nominotypical) (Japanese) mature (IV—V) immature (II) 129-202 177-302 182* 150.26 + 0.73 224.85 + 4.49 147-315 209-390 184.23 + 0.78 273.48 + 0.91 50-129 58-202 no data 1822 += 055 107.13 + 0.45 0.485—0.523 0.445-0.512 0.501 + 0.006 0.486 + 0.004 (0.500)* 0.687—0.827 0.643-0.739 0.771 + 0.018 0.695 + 0.005 0.643* weak pronounced weak* 1.6—2.0 2.0—2.8 )R,-L, M1, M2, iR\-L; M1, M2, no data dicuspid tricuspid 3755 25,000 no data 5.94 4.20 no data 0.3—0.5 3.0—4.0 no data quasibenthic quasibenthic stenobathic stenobenthic eurybathic(?) spring summer, autumn no data * Holotype (Okutani et al., 1987). RO-Centre), Vladivostok, Russia. Holotype: BMJSO01. Paratype: BMJSO02. Type locality: Type specimens were collected by Pavel V. Kaltchugin on the Russian research vessel Professor Kaganovskyi from the continental slope of the Japan Sea (Russian Primorie region, 42°30'N, 133°42'E); bottom trawling station; 600 m deep; 23 May 1996. Diagnosis: Subspecies small (fully mature females usu- ally to 200 mm DML, mature males usually to 170 mm DML); fin long (to 50% of DML) and wide (to 77% of DML); poorly shaped, and weakly differentiated hookless tentacular club; small differences between central and marginal suckers on the manus of the club; fixing appa- ratus undifferentiated, consists of almost 40 elements (42 pads alternating with minute hookless suckers in holo- type), starts from approximately central dorsal part of the stalk, and terminates on distal part of the manus; bicuspid lateral teeth of the radula. Description: Body (Figure 8). Small to medium size. Dorsal mantle length (DML) of mature prespawning fe- males usually does not exceed 180-200 mm, Mature males usually do not exceed 140-170 mm. Mantle. Cylindrical; width (MW) approximately 1/3 of DML: MW 36% of DML for holotype. Anterodorsal mar- gin of mantle with triangular projection; small in imma- ture adults, well-defined in prespawning and spawning animals. Anteroventral mantle edge slightly emerginate with two small lateral angles which border funnel. Fin. Rhomboid, large; length (FL) about 50% of DML, width (FW) almost 80% of DML. Anterior and posterior fin margins straight; anterior lobes of fin distinct; lateral margins slightly rounded. Head. Cubic and large; slightly smaller than mantle opening. Head width index (HWI) 25% in holotype. Eyes large, with long, deep sinus; eye diameter index (EDI) about 14% in holotype. Funnel organ (Figure 9). Composed of dorsal element (DFO) and two smaller ventral elements (VFO). Dorsal pad large, A-shaped. Both limbs of DFO joined anteri- O. N. Katugin, 2000 AIV (HC) AIV (FA) Figure 8 Berryteuthis magister shevtsovi Katugin, subsp. nov. Dorsal view of the holotype. DDVV, type of buccal lappets attachment (Dor- sal, Dorsal, Ventral, Ventral); AI—AIV, arms one to four; HC, hectocotylized arm; FA, fellow arm; DML, dorsal mantle length; MW, mantle width; HW, head width; ED, eye diameter; FL, fin length; FW, fin width. orly; apex bifurcal. Total length of DFO almost 15% of DML in holotype. Ventral pads oval, 2/3 length of DFO. Mantle locking apparatus (Figure 9). Consists of nu- chal cartilage (NC), and funnel cartilages (FC). Nuchal cartilage elongated, slightly broader proximally, with me- dian and two lateral grooves; length nearly 12% of DML. Funnel locking-cartilage lanceolate with median sulcus anteriorly; flat and expanded posteriorly; length almost 14% of DML in holotype. Mantle cartilages (MC) which correspond to funnel cartilages consist of low ridges which expand posteriorly. Arms (Figure 10A, B). Muscular and long; longest arm 70%, and shortest 61% of DML in holotype. Arm for- Page 93 (A) MLC (B) Figure 9 Berryteuthis magister shevtsovi Katugin, subsp. nov. Head, lock- ing cartilages, and funnel organ. A. Ventral view (mantle is opened). MLC, mantle locking cartilage; FLC, funnel locking cartilage; DFO, dorsal funnel organ (inside the funnel); VFO, ventral funnel organ (inside the funnel). B. Nuchal cartilage (NC). mula typical for species: II/>II>I>IV. Aboral swimming keel well developed on third arm, less evident on other arms. Arms equipped with tetraserial armature. Armature in four longitudinal rows (ventral, medioventral, medi- odorsal, and dorsal). Basal or proximal regions of all arms lack armature; first suckers appear on ventral and dorsal rows; distal parts of all arms with minute suckers alone. Ventral arms (AIV) with suckers only. In adult males cen- tral or medial region of one of arm IV pair modified as hectocotylus (HC); opposite ventral or fellow arm (FA) not modified. Females lack modifications on arms IV. Arms I, II, and III bear suckers in ventral and dorsal rows and hooks in central regions of medioventral and medi- odorsal rows. In each central row five to six suckers pre- sent, followed by 11 to 13 hooks. Number of hooks on arms I, II, and III in holotype 22, 24, and 26, respectively. Overall numbers of transverse rows of armature on arms I-IV in holotype 31, 32, 37, and about 40, respectively. Hectocotylus. Modifications confined to medial region of one ventral arm only; proximal third and distal third of arm appeared unmodified (Figure 10A, B). Suckers in dorsal and mediodorsal rows differ in size and form, while suckers in medioventral and ventral rows remain unchanged. Modified sucker stalks of both rows suffi- Page 94 1 mm 1 mm The Veliger, Vol. 43, No. 1 AI-III AIV (HC) °° >i -5-) ° ® ° ° 4° 2, Bo ° ho 25 ae 2, %, 5 . hoo 4 wr) 20 < Po %, Co) ° o role Ps e, o 3° ° ° ° QO SC SCOg Figure 10 Berryteuthis magister shevtsovi Katugin, subsp. nov. A. Hectocotylized ventral arm of the holotype specimen. AIV(HC), ventral view of the arm; NP, unmodified sucker (enlarged); NSR, unmodified sucker ring (enlarged); MP, modified sucker (enlarged); DSR, modified sucker ring (enlarged). B. Diagram of arm measurements: AI-III, arms one to three; AITV(HC), arm four (hectocotylized arm); AL, arm length; A, basal region without suckers; B, proximal suckers zone; C, hooks zone; D, distal suckers zone; HCL, zone with modified suckers. ciently enlarged to form narrow groove. Sucker modifi- cations in fully mature and prespawning males much more conspicuous than in immature and maturating spec- imens. Left ventral arm modified in holotype and two specimens; in all others right ventral arm modified. Hec- tocotylus of holotype described below. First eight suckers in dorsal row, and first six suckers in mediodorsal row appear almost unmodified, except some thickening at base of sixth and eighth sucker stalks. Noticeable changes evident in 11 succeeding suckers (rows 9-19) in dorsal row, and in ten succeeding suckers (rows 7-16) in me- diodorsal row. Papilla formed by stalk base of each mod- ified sucker, 5-6 mm high; sucker pedicel (stalk upper part) short, about | mm in length; sucker ring almost two times smaller than that of unmodified sucker. Number of unmodified proximal, and affected central suckers varies among specimens, six to nine and eight to twelve in dor- sal row; six to eight and six to nine in mediodorsal row. Ventral arm modifications in B. magister were first de- scribed as a true hectocotylus in several mature males from the northeastern Pacific, and Bering Sea by Voight (1995). Such arm modifications in males of B. magister from the Japan Sea are herein reported for the first time. Holotype and ten males of squid in various maturity stag- es (II-V) were analyzed. All specimens, except one im- mature (stage II) male, had noticeable morphological changes on one of the ventral arms. Tentacles (Figure 11A, B). Long, narrow, almost 1.5 times longer than mantle length. Tentacle total length (TTL) of holotype 157% of DML. Club elongated, lan- ceolate; maximum width (CLMW) 4% of TTL. All club regions (e.g., carpal zone, manus, and dactylus) weakly defined (Figure 11A). Principal measurements made based on attachment of club membranes (Figure 11B). Boundary between dactylus and manus roughly corre- sponds to beginning of apical aboral membrane on outer surface of club; relative length of dactylus (CLWL/TTL) 8.2% in holotype. Proximal boundary of manus corre- sponds to starting point of ventral membrane; length of this membrane (CLVL) 20% of TTL in holotype; ratio of manus length to TTL 11.8%. Proximal attachment of dor- sal membrane corresponds to boundary between club and stalk; length of this membrane (CLDL), and thus club length, 28% of TTL, and 44% of DML in holotype. Car- O. N. Katugin, 2000 Dactylus Manus Page 95 CLMW Figure 11 Berryteuthis magister shevtsovi Katugin, subsp. nov. A. Tentacle of the holotype specimen with details of the club (carpus, manus, and dactylus). S, distal part of the stalk; ACS, apical circlet of suckers; AW, aboral web; DM, dorsal membrane; FA, fixing apparatus (enlarged are two pads, and two alternating toothless suckers); VM, ventral membrane. B. Diagram of tentacle measurements: TTL, total tentacle length; SZL, suckers zone length; CLDL, club dorsal length (from the place of dorsal membrane attachment); CLVL, club ventral length (from the place of ventral membrane attachment); CLWL, club aboral web length; CLMW, club maximum width. pal zone covers about 8% of holotype TTL, and some- what less, than 30% of club length. Sucker zone length (SZL) occupies almost 46% of TTL in holotype. Circlet of minute suckers at tip of tentacle (Figure 11A); ten suckers on right tentacle, and 11 suckers on the left; circlet diameter 2.4 mm in holotype. Sucker rim without teeth; diameter of sucker rim ranges from 0.3— 0.6 mm. Proximal to apical circle transverse rows of suckers present on inner surface of dactylus. Two distal- most sucker rows tetraserial; number of suckers in each transverse row increases proximally. Suckers toothless to central part of dactylus where bluntly pointed teeth first appear. Largest suckers in central part of manus; maxi- mum ring diameter 0.6—0.75 mm. Small suckers cover dactylus, borders of manus, carpal zone, and proximal stalk; minimum diameter of rim of sucker from ventral margin of manus 0.3—0.32 in holotype; ratio of largest to smallest rim diameter range from 1.6—2.0 in holotype. Fixing apparatus (Figure 11A). Extends to dorsal edge of tentacle, starting from first suckers on stalk, and ter- minating on border between manus and dactylus. Locking zone consists of alternating series of pads (or knobs) and minute toothless suckers. Knob diameter 1.5—2 times larger than sucker ring diameter; number of knobs ap- proximately 40 in holotype. Distribution and habitat: The range of B. (m.) shevtsovi is presumably limited to the Japan Sea, and data on dis- tribution and ecology of B. magister in this region could be applied to the new subspecies as well. This squid has been reported between the Oki and Tsushima groups of islands (35°25'N, 130°35’E, E. V. Slobodskoy, personal communication), hence the southernmost boundary of distribution presumably goes through the Korean Strait. It is widely distributed throughout the Japan Sea, with large stocks reported from the Jamato Bank, Kitayamato Page 96 The Veliger, Vol. 43, No. 1 Bank, Sinoki Bank, Rebun Bank, and from the Oki and Noto Seamounts (Ogata et al., 1973; Kasahara et al., 1978; Nazumi et al., 1979; Natsukari et al., 1993; Shev- tsov, 1988). The northernmost boundary is in the Tatarsky Strait (Railko, 1979). Vertical distribution of the squid in the Japan Sea is uneven. Adults are confined to the slope depths. Specimens of various maturity stages ranged in depth from 50—1200 m with a maximum occurrence of adults in depths 300-500 m, which is characteristic of the species (Kubodera, 1982, 1992; Okiyama, 1993; Railko, 1979). Habitat is determined by rather specific hydrolog- ical conditions of the Japan Sea, when compared to the subarctic hydrology of the rest of the northern Pacific Ocean (Okiyama, 1993; Kitano, 1958). The most notable feature of the squid’s habitat in the Japan Sea is low tem- perature, in the range of 0.2—1.5°C (Okiyama, 1993; Rail- ko, 1979). Etymology: In honor of Dr. Gennadyi A. Shevtsov, em- inent Russian researcher for the North Pacific cephalo- pods, particularly for the cephalopods from the Japan Sea. Comparative morphologic, reproductive, and ecologic features for the three subspecies of B. magister, namely B. (m.) magister;, B. (m.) nipponensis; and B. (m.) shev- tsovi are presented in the Table 7. Taxonomic summary: B. (m.) shevtsovi is distinguish- able both from B. (m.) magister, and from B. (m.) nip- ponensis, though as expected, morphological intersubspe- cific differences are rather quantitative. There are virtu- ally no diagnostic morphologic characters, which could unambiguously (with a 100% probability of correct di- agnosis) distinguish among all the three subspecies. Ma- ture adults of B. (m.) shevtsovi are considerably smaller, have a comparatively larger fin, and less pronounced dif- ferences between central and marginal club suckers than mature specimens of B. (m.) magister. Most specimens of the new subspecies possess bicuspid lateral teeth of the radula, while there are three cusps on the same teeth in the majority of specimens of the nominotypical sub- species from the Bering Sea. It is rather hard to compare taxonomic characters of B. (m.) shevtsovi with those of B. (m.) nipponensis, because the latter was described from immature specimens with undeveloped traits of external morphology. We can guess though, that basic differences between these taxa are in body size (the new subspecies is considerably smaller), in fin size (the new subspecies has a larger fin), and in mantle width (the new subspecies has a wider mantle with pallial aperture larger than head width). All the three subspecies also differ in ecological characters, B. (m.) magister being stenobathic, B. (m.) shevtsovi—cold-water stenobathic, and B. (m.) nipponen- sis—presumably eurybathic. Acknowledgments. I thank Pavel V. Kaltchugin for collecting specimens for taxonomic description of the new subspecies of B. magister, Vyacheslav G. Rybin for help in computer presentation of data on squid morphology, Peter P. Railko for help in analyz- ing data on morphologic variation of squid, Dr. Gennadyi A. Shevtsov for providing biological data on squid and useful com- ments on my manuscript, and especially Dr. E G. Hochberg for his encouragement and priceless help during preparation of this paper. 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Genetic evidence of population heterogeneity and cryptic speciation in the ommastrephid squid Martialia hy- adesi from the Patagonian shelf and Antarctic Polar Frontal Zone. Marine Biology 116(4):593-602. BRIERLEY, A. S., J. P. THORPE, M. R. CLARKE & H. R. MARTINS. 1993b. A preliminary biochemical genetic investigation of pop- ulation structure of Loligo forbesi Steenstrup, 1856 from the British Isles and the Azores. Pp. 61—69 in T. Okutani, R. K. O’Dor & T. Kubodera (eds.), Recent Advances in Cephalopod Fisheries Biology. Tokai University Press: Tokyo. CARVALHO, G. R. & T. J. PITCHER. 1989. Biochemical genetic studies on the Patagonian squid Loligo gahi d’Orbigny. II. Population structure in Falkland waters using isozymes, morphometrics and life history data. Journal of Experimen- tal Marine Biology and Ecology 126(3):243-258. CARVALHO, G. R., A. THOMPSON & A. L. STONER. 1992. Genetic diversity and population differentiation of the short-fin squid Illex argentinus in the southwest Atlantic. Journal of Exper- imental Marine Biology and Ecology 158:105—121. ELDREDGE, N. & S. J. GouLp. 1972. Punctuated equilibria: an alternative to phyletic gradualism. Pp. 82-115 in T. J. M. Schopf (ed.), Models in Paleobiology. Freeman, Cooper: San Francisco. Feporets, YU. A. 1983. Seasonal distribution of the squid Ber- ryteuthis magister in the western Bering Sea. Pp. 129-130 in Systematics and Ecology of Cephalopods. Zoological In- stitute of Academy of Sciences of the USSR: Leningrad. [in Russian] Feporets, Yu. A. & O. A. KozLova. 1986. Reproduction, fe- cundity, and abundance of the squid Berryteuthis magister (Gonatidae) in the Bering Sea. Pp. 66-80 in Resources and the Outlook for Utilization of Squid in the World Ocean. VNIRO: Moscow. [In Russian] KASAHARA, S., T. NAZUMI, T. SHImMIzuU & H. HAMABE. 1978. Con- tributions of biological information useful for development of inshore squid fishery in the Japan Sea. II. A note on reproduction and distribution of Berryteuthis magister (Ber- ry) assumed from biological observations on trawl catches in the waters around the Oki Islands, Japan Sea. Bulletin of O. N. Katugin, 2000 the Japan Sea Regional Fisheries Research Laboratory 29: 159-178. [in Japanese with English summary] Katuain, O. N. 1991. Unidentified gangle protein polymorphism in the gonatid squid Berryteuthis magister from the North Pacific. Isozyme Bulletin 24:51. KatuaIn, O. N. 1992. Polymorphism of purinenucleoside phos- phorylase in the North Pacific oceanic squids. Isozyme Bul- letin 25:56. Katuain, O. N. 1993. Genetic variation in the squid Berryteuthis magister (Berry, 1913) (Oegopsida: Gonatidae). Pp. 201— 213 in T. Okutani, R. K. O’Dor & T. Kubodera (eds.), Recent Advances in Cephalopod Fisheries Biology. Tokai Univer- sity Press: Tokyo. KatuGciIn, O. N. 1995. 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American Journal of Human Genetics 22:24—29. WRIGHT, S. 1978. Evolution and the Genetics of Populations. Vol. 4. Variability Within and Among Natural Populations. University of Chicago Press: Chicago. YuukI, Y. & H. KITAZAWA. 1986. Berryteuthis magister in the southwestern Japan Sea. Bulletin of the Japanese Society of Scientific Fisheries 52(4):665—672. ZAKHVATKIN, YU. A. 1975. Embryology of Insects. ‘““Wyshaya Shkola’’: Moscow. [in Russian] The Veliger 43(1):98-102 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 NOTES, INFORMATION & NEWS Arboreal Neritidae Robert H. Cowie! and Barry D. Smith? Neritidae are snails of marine, brackish, and freshwater aquatic environments. This note reports a species Neri- todryas cornea (Linnaeus, 1758), living in trees on the island of Babeldaob in the Palau archipelago of the west- ern Pacific. This observation is noteworthy because there are few other explicit reports in the literature of neritids living in trees, an apparently unusual habitat for the fam- ily. Neritodryas cornea were observed and collected at a site in the southwest part of Babeldaob (Imeliik State) (134°30.7'E, 7°24.2'N) on 5 June 1996. The site is a grove of pandanus trees, Pandanus cf. kanehirae Martelli in Kanehira, 1933, growing between the inland boundary of a mangrove forest and the unpaved road that travels north-south in this part of the island, approximately 6.5 km north of the Koror-Babeldaob (“KB”) Bridge (now collapsed). A small stream runs under the road, through the pandanus grove and into the mangroves. The ground under the pandanus was waterlogged. No Neritodryas cornea were found on the ground at this site, or in the adjacent mangroves or stream. The snails were found in the pandanus trees, predominantly at the bases of the leaves where they joined the stem of the tree and where small amounts of water collected, but also on other parts of the upper surfaces of the leaves. They were found in some of the tallest pandanus trees, up to 3 m above ground. There was no evidence that the snails were in the trees by accident, as a result of being stranded after receding floods. The area clearly does not flood to the height at which the snails were found, and there had been no re- cent storm activity. Thus we consider the snails’ pres- ence at these heights in the pandanus to be normal. Whether they move up and down the trees is not known, or whether and how they move between trees. Whether they have a short-lived larval stage or none, as might perhaps be appropriate for their arboreal habit, is also not known. It is known that some neritid species, including Neri- todryas cornea, are frequently found above water. Indeed N. cornea has been collected previously (but not reported) from the same arboreal habitat in Palau (B. Holthuis, per- sonal communication), and from similar habitat in New ' Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817, USA. ? Marine Laboratory, UOG Station, University of Guam, Man- gilao, Guam 96923, USA. Guinea (M. Pouliceck, personal communication). And other species in the genus are usually somewhat terrestrial (e.g., Baker, 1923), found on vegetation in the vicinity of water (Franc, 1956; B. Holthuis, personal communica- tion; C. Unabia, personal communication). Also on Ba- beldaob, Smith (unpublished report to The Nature Con- servancy, University of Guam Marine Laboratory, 1991) reported Neritodryas subsulcata (Sowerby, 1836) up to 2 m above the waterline on river banks. In Papua New Guinea, Nerita planospira Anton, 1839, has been found in pandanus and other trees in the mangrove forest (L. M. Cook, personal communication). Neritodryas cornea is a widely distributed species, re- ported not only from Palau but also from New Guinea (van Benthem Jutting, 1963; Pouliceck et al., 1994); Sol- omon Islands (Haynes, 1990, 1993); and from the Nico- bar Islands, through Java and the Malay archipelago to the Philippines and New Caledonia (van Benthem Jutting, 1956). Given the many misidentifications of neritids in the literature, it could in reality be more (or less) widely distributed. Other neritids recorded in the collection area were Ner- itina turrita (Gmelin, 1791) on the muddy substrate at the edge of the mangroves immediately adjacent to the pan- danus grove, but not in the grove itself, and Neritina squamaepicta (Récluz, 1843) in the stream as it flowed into the grove. All specimens are deposited in the Malacology collec- tions of Bishop Museum (Honolulu): catalog numbers BPBM 253989 (Neritodryas cornea), BPBM 253990 (Neritina turrita), BPBM 253992 (Neritina squamaepic- ta). Acknowledgments. The snails were discovered in the pandanus by Allen Allison and Chris Austin. We thank Lynn Raulerson and Derral Herbst for identifying the pandanus, and Bern Hol- thuis and Anne Brasher for discussion. We also thank the follow- ing additional people for responding to our request for infor- mation about neritids in trees: Laurence Cook, James Carucci, Mathieu Pouliceck, and Catherine Unabia. Literature Cited Baker, H. B. 1923. Notes on the radula of the Neritidae. Pro- ceedings of the Academy of Natural Sciences of Philadel- phia 75:117-178, pls. 9-15. FrAnc, A. 1956. Mollusques terrestres et fluviatiles de l’archipel Néo-Calédonien. Mémoires du Muséum National d’ Histoire Naturelle, Série A. Zoologie 13:1—200, pls. 1-25. Haynes, A. 1990. The numbers of freshwater gastropods on Pa- cific islands and the theory of island biogeography. Mala- cologia 31(2):237-248. Haynes, A. 1993. The gastropods in the streams and rivers of Notes, Information & News Page 99 four islands (Guadalcanal, Makira, Malaita, and New Geor- gia) in the Solomon Islands. The Veliger 36(3):285—290. Pou.iceck, M., J. C. Bussers & P. VANDEWALLE. 1994. Ecologie des Neritidae et autres Gastéropodes intertidaux de l’ile de Laing (Papouasie Nouvelle-Guinée). Haliotis 23:15-31. VAN BENTHEM JuTTING, W. S. S. 1956. Systematic studies on the non-marine Mollusca of the Indo-Australian Archipelago. V. Critical revision of the Javanese freshwater gastropods. Treubia 23(2):259—477. VAN BENTHEM JUTTING, W. S. S. 1963. Non-marine Mollusca of west New Guinea. Part 1, Mollusca from fresh and brackish waters. Nova Guinea, Zoology 20:409-S21. Paleogeographic Implications of Late Paleocene Onestia onestae (Bivalvia: Cardiidae) in Arctic Alaska Jay A. Schneider! and Louie Marincovich, Jr. Introduction The northernmost and youngest occurrence of the cardiid bivalve Onestia onestae (McLearn, 1931) is reported from Upper Paleocene beds (Marincovich, 1996; Bice et al., 1996) in the Prince Creek Formation near Ocean Point, northern Alaska (70°04’N, 151°22'W). The only previous occurrences of O. onestae are in middle Creta- ceous strata of central Alberta, Canada: the upper Aptian Clearwater Shale (McLearn, 1933) (approx. 57°N, 112°W) and the lower Albian lower sandstone member of the Peace River Formation (McLearn, 1931) (approx. 56°N, 117°W). This is the only occurrence of Onestia in the northern hemisphere. Three other species of Onestia occur in Aptian and Albian faunas of Australia (Day, 1978). Systematic Paleontology Family CARDIDAE Lamarck, 1809 Subfamily LAHILLINAE Finlay & Marwick, 1937 Genus Onestia McLearn, 1933 Onestia onestae (McLearn, 1931) (Figures 1-7) Laevicardium onestae McLearn, 1931:7, pl. 1, fig. 1. Integricardium (Onestia) onestae (McLearn) McLearn, 1933:152-153, pl. 2, figs. 8-10. Onestia onestae (McLearn) McLearn, 1945:10, pl. 3, fig. 9; ' Department of Geology and Geophysics, University of Wis- consin, 1215 W. Dayton Street, Madison, Wisconsin 53706, USA. 2 Department of Invertebrate Zoology and Geology, California Academy of Sciences, San Francisco, California 94118, USA. Table 1 Dimensions of complete valves of Onestia onestae. See Schneider (1995, in press) for explanations of height, length, and anterior length. Dimensions (mm) Anterior Height Length length Inflation GSC 6345 24.7 31.6 10.1 9.1 GSC 8004 26.0 31.0 13.0 10.0 USGS [M9158] (right valve) 16.4 18.9 Hail 5.3 USGS [M8120] (left valve) 26.2 31.9 12.5 OS) USGS [M8120] (right valve) 27.6 30.7 14.4 9.4 Jeletzky, 1964:9, 76, pl. 24, figs. 9, 11; Day, 1978:38, pl. 1, figs. 1-3. Description: Elliptical in shape, moderately inflated, um- bones located slightly anteriorly (see Table 1 for mea- surements). Anterior and ventral margins convex, poste- rior margin less convex and slightly oblique. Sculpture consists of growth lines only. Beaks low, slightly proso- gyrous. Ligament groove short. Pallial sinus absent. Ad- ductor muscles subcircular, faintly impressed. Hinge teeth arranged as in Protocardia and Integricardium (see Schneider, 1995), but anterior cardinals more robust. Right anterior cardinal thick and blunt, right posterior car- dinal strong, subconical, and pointed. Right anterior car- dinal socket deep and conical. Right posterior cardinal socket shallower and usually broader. Right anterior lat- eral tooth weak; overlying anterior lateral socket shallow. Right posterior lateral long, narrow, but not bladelike. Overlying posterior lateral socket moderately deep. No known specimens of left valves are well-preserved enough to describe hinge in detail. Material: Holotype, Geological Survey of Canada-GSC 6345, left valve (Figure 1), lower Albian Peace River Formation, Alberta; plesiotypes GSC 8003 (left valve) and 8004 (right valve, Figure 2), upper Aptian Clearwater Shale, Alberta; University of California Museum of Pa- leontology, Berkeley, California. UCMP 154061 (right valve, Figures 3 and 6) and UCMP 154062 (left valve, Figures 4, 5), United States Geological Survey, USGS locality M8120, Ocean Point, Alaska; UCMP 154063 (Figure 7) and 154064 (two right valves), USGS locality M9158, Ocean Point, Alaska; UCMP 154065, 154066, and 154067 (three right valves), USGS locality M8120; all UCMP specimens from Upper Paleocene Ocean Point beds of Prince Creek Formation. Discussion There is a stratigraphic gap of some 45 to 50 million years (Harland et al., 1990) between the Canadian and Alaskan occurrences of O. onestae, but the Paleocene Page 100 The Veliger, Vol. 43, No. 1 Explanation of Figures | to 5 Figure 1. Onestia onestae, external view of left valve, X 1.35, GSC 6345. Figure 2. Onestia onestae, internal view of right valve, * 1.35, GSC 8004. Figure 3. Onestia onestae, external view of right valve, X 1.35, UCMP 154061. Figure 4. Onestia onestae, internal view of left valve, 1.35, UCMP 154062. Figure 5. Onestia onestae, external view of left valve, X 1.35, UCMP 154062. specimens are so similar to the Cretaceous specimens that erection of a new species for the former is unwarranted. The similarity between the Aptian-Albian and Paleocene specimens may partly result from Onestia having lost or reduced the characters that are normally used to discern cardiid species: the external ornamentation of most car- diids is lost, and the lateral teeth are reduced. Shell shape is most commonly used to discriminate between species of Cardiidae which have undergone such losses and re- duction, such as within Onestia’s fellow lahilliines Lahil- lia (Cossmann, 1899) and Jntegricardium (Rollier, 1912); and the Canadian and Alaskan forms of Onestia cannot be discriminated on the basis of shell shape (Table 1). The Paleocene forms are therefore classified as Onestia onestae. Marincovich (1993) reported two additional cardiids from the Ocean Point beds: Integricardium (UIntegricar- dium) keenae Marincovich, 1993, and Protocardia? sp. indet. One other species of Integricardium occurs in North America, /ntegricardium holmesi (Russell, 1943), from the Maastrichtian Eastend Formation of Saskatche- wan, but the genus is otherwise known only from Het- tangian to Maastrichtian-age rocks in Africa and Eurasia (Keen, 1969; Marincovich, 1993; Schneider, 1995). Al- though the generic assignment of Protocardia? sp. indet. by Marincovich (1993) is uncertain, Protocardia is known from all continents except Australia and Antarc- tica, and has a stratigraphic range of Rhaetian to Maas- trichtian (Keen, 1969; Schneider, 1995). Onestia and Integricardium are two of six genus-level bivalve taxa in the Ocean Point beds that otherwise have their youngest occurrences in Mesozoic faunas (Marin- covich, 1993). Camptochlamys Arkell, 1930, is otherwise known only from the Late Jurassic. The next youngest occurrence of Tancredia (Tancredia) Lycette, 1850, is Albian. The next youngest occurrences of Oxytoma (Hy- poxytoma) Ichikawa, 1958, and Tellinimera Conrad, 1860, are Maastrichtian. These six of the 24 genera re- ported from the Ocean Point beds are considered to be relict taxa. These six genera were diverse and widely dis- tributed during the Mesozoic, but by the Late Paleocene they were represented by a single species living only in the Arctic Ocean (Marincovich, 1993, 1996). However, Onestia is the only one of these six taxa to be represented by a species known from the Cretaceous (Marincovich, 1993). Although detailed systematic analyses of these rel- ict bivalve taxa remain to be done, the Paleocene species of these taxa do not appear to be as evolutionarily con- servative as Onestia onestae. The presence of Onestia onestae in Upper Paleocene strata of northern Alaska reinforces the concept that relict Mesozoic mollusks lived into Cenozoic time within a geographically isolated Arctic Ocean (Marincovich, 1993, 1996). The occurrence of O. onestae in northern Alaska Notes, Information & News Page 101 6 5mm 10mm Explanation of Figures 6 and 7 Figure 6. Onestia onestae, detail of hinge of right valve, UCMP 154061. Figure 7. Onestia onestae, interior view of right valve showing adductor muscles and pallial line, UCMP 154063. supports the idea that a significant portion of Paleocene mollusks in the Arctic realm descended from Western In- terior Seaway Cretaceous faunas. This find extends the geographic range of Onestia into high northern latitudes where it was previously unknown. Acknowledgments. Type material of Onestia onestae was pro- vided by T. E. Bolton of the Geological Survey of Canada, and comparative specimens of /ntegricardium holmesi were provided by J. Waddington of the Royal Ontario Museum. Literature Cited Bice, K. L., M. A. ARTHUR & L. MARINCOVICH, JR. 1996. Late Paleocene Arctic Ocean shallow-marine temperatures from mollusc stable isotopes. Paleoceanography 11:241—249. Day, R. W. 1978. Onestia McLearn, an unusual cardiacean pe- lecypod from the Lower Cretaceous of Australia and Can- ada. Department of National Development, Bureau of Min- eral Resources, Geology and Geophysics, Bulletin 192:37— 44. HARLAND, W. B., R. L. ARMSTRONG, A. V. Cox, L. E. CRAIG, A. G. SmitH & D. G. SmitH. 1990. A Geologic Time Scale 1989. Cambridge University Press: New York. 263 pp. JELETZKY, J. A. 1964. Illustrations of Canadian fossils. Lower Cretaceous marine index fossils of the sedimentary basins of western and Arctic Canada. Geological Survey of Canada Paper 64-11. KEEN, A. M. 1969. Superfamily Cardiacea Lamarck, 1809. Pp. 583-594 in R. C. Moore (ed.), Treatise on Invertebrate Pa- leontology, part N, volume 2, Mollusca 6, Bivalvia. The Geological Society of America and the University of Kansas Press: Lawrence, Kansas. Marincovicu, L. JR. 1993. Danian mollusks from the Prince Creek Formation, Northern Alaska, and implications for Arctic Ocean paleogeography. Paleontological Society Memoir 35:1-35. Marincovicu, L. JR. 1996. Survivorship of Mesozoic mollusks in the Paleocene Arctic Ocean. Pp. 275-288 in N. MacLeod & G. Keller (eds.), The Cretaceous-Tertiary Mass Extinc- tion: Biotic and Environmental Changes, W. W. Norton: New York. McLEArRN, F. H. 1931. The Gastroplites and other Lower Creta- ceous faunas of the Northern Great Plains. Transactions of the Royal Society of Canada, section 4, series 3, 25:1—7. McLEARN, FE H. 1933. Pelecypods of the Lower Cretaceous Clearwater Formation, Northern Alberta. Transactions of the Royal Society of Canada, section 4, series 3, 27:139—-156. McLearn, E H. 1945. Revision of the Lower Cretaceous of the western interior of Canada, 2nd ed. Geological Survey of Canada Paper 44-17:1—14. RUSSELL, L. S. 1943. Marine fauna of the Eastend Formation of Saskatchewan. Journal of Paleontology 17:281—288. SCHNEIDER, J. A. 1995. Phylogeny of the Cardiidae (Mollusca: Bivalvia): Protocardiinae, Laevicardiinae, Lahilliinae, Tu- longocardiinae subfam. n. and Pleuriocardiinae subfam. n. Zoologica Scripta 24:321-346. SCHNEIDER, J. A. In press. Phylogeny of stem-group ‘‘eucardiids”’ (Bivalvia: Cardiidae) and the significance of the transitional fossil Perucardia. Malacologia. Page 102 International Commission on Zoological Nomenclature The new and extensively revised 4th Edition of the Jn- ternational Code of Zoological Nomenclature has now been published and is in effect from 1 January 2000. The price is US $65 or £40, but discounts are offered to in- dividuals buying the Code for personal use or to institu- tions buying five or more copies. Full details of how to buy copies are given on the Commission’s website (Www.iczn.org) or may be obtained by e-mailing ““iczn@nhm.ac.uk.”’ The following Application was published on 30 June 1999 in Volume 56, Part 2 of the Bulletin of Zoological Nomenclature. Comment or advice on this application is invited for publication in the Bulletin and should be sent to the Executive Secretary, I.C.Z.N., c/o The Natural His- tory Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). The Veliger, Vol. 43, No. 1 Case 3126—Bulinus wrighti Mandahl-Barth, 1965 (Mol- lusca, Gastropoda): proposed conservation of the spe- cific name. The following Opinions concerning mollusks were pub- lished on 30 June 1999 in Volume 56, Part 2 of the Bul- letin of Zoological Nomenclature. Copies of this Opinion can be obtained free of charge from the Executive Sec- retary at the address given above. Opinion 1924. Helix draparnaudi Beck, 1837 (currently Oxychilus draparnaudi; Mollusca, Gastropoda): specif- ic name conserved. Opinion 1925. Turrilites gravesianus d’Orbigny, 1842 (currently Hypoturrilites gravesianus; Mollusca, Am- monoidea): specific name conserved and a replacement lectotype designated; Turrilites tuberculatus Bosc, {1802] (currently Hypoturrilites tuberculatus): placed on the Official List. The Veliger 43(1):103—104 (January 3, 2000) THE VELIGER © CMS, Inc., 2000 BOOKS, PERIODICALS & PAMPHLETS Bivalves: An Eon of Evolution. Paleobiological Studies Honoring Norman D. Newell edited by P. A. JOHNSTON AND J. W. HAGGART. 1998. Uni- versity of Calgary Press, 2500 University Drive N. W., Calgary, Alberta, Canada T2N 1N4. 461 pp. This volume, dedicated to Norman D. Newell, is at once a fitting tribute to one of the most diverse bivalve paleontologists and systematists of this century, and a misnomer. Those interested in a synthesis of current knowledge concerning the evolution of the Bivalvia will be disappointed. The volume instead is a potpourri of papers encompassing ecology, paleoecology, functional morphology, taphonomy, taxonomy, and studies of the most ancient relationships among bivalved mollusks. This diversity of papers is indeed a tribute to the breadth of Newell’s contributions to bivalve paleobiology. The most significant and influential paper of the vol- ume is the lead article by Waller (‘Origin of the mollus- can Class Bivalvia and a phylogeny of major groups’’). Waller tackles some of the most vexing and lingering problems in malacology, namely the origin of the phylum and the relationships among the major classes. The sheer amount of character information and analysis presented in this paper is truly impressive. Waller presents a series of nested and well-supported hypotheses beginning with the relationships of the Bivalvia to the other molluscan classes, and proceeding to analyses of major bivalve or- ders. The paper, to my knowledge, is to date the most informed and informative analysis on this topic. It does, however, highlight a common and problematic theme throughout the volume, and that is the underutilization of modern phylogenetic concepts and methodology. While few papers in the volume are actually dedicated to phylogenetic analyses, a number of them address topics that require phylogenetic information. For example, Col- lom’s report on the taxonomy and biostratigraphy of Cremnoceramus hinges upon assumptions of ancestry in the Late Turonian. It would be nice to see these assump- tions subjected to a phylogenetic analysis, as it would to have a character data set reconstructed from Waller’s re- port and subjected to a parsimony analysis. On the other hand, Collom’s paper is a gem regarding the incorpora- tion of ontogenetic and paleoecological information into a biostratigraphic and taxonomic study. Likewise, one would be hard pressed to match the depth of understand- ing that Waller possesses concerning the fundamental characters of bivalve evolution and diversity. Examples of this contrast abound in the volume. Bringing all these varied fields and types of information together stands to- day as one of the major challenges facing paleontologists and others seeking to understand the evolution of organ- isms with fossil records as excellent as the Bivalvia’s. There are other contributions that are dedicated pri- marily to phylogenetic analysis, or rely upon the results of such analyses. The most ambitious of these is Camp- bell et al.’s “18S ribosomal DNA and evolutionary rela- tionships within the Bivalvia,” an analysis of relation- ships among major bivalve superfamilies. While the num- ber of base pairs reported (800) is rather low, and the analyses were restricted to heuristic searches because of the tremendous size of the data set, this is currently the most promising project for elucidating higher relation- ships within the Bivalvia. At lower levels, papers by Frischer et al. (‘A molecular phylogeny of some major groups of Pectinidae inferred from 18sRNA gene se- quences’’) and Carter & Seed (“‘Thermal potentiation and mineralogical evolution in Mytilus (Mollusca; Bivalvia)) reflect the breadth of approaches that can be used to un- derstand bivalve systematics. The other major foci of the volume are the paleoecol- ogy and biostratigraphy of bivalves. Bivalves are critical to both these disciplines during various intervals because of their excellent fossil record. Their numerical promi- nence and diversity in many fossil assemblages and fau- nas since the Silurian, coupled with the general applica- bility of uniformitarian interpretations to understanding their paleoecologies, has rendered bivalves one of the most utilized groups in paleoecological research. Much of Newell’s work stands as testament to this. Several pa- pers in the volume highlight this fact, including the ex- tensive interpretive compilation of reef-associates by Eliuk. A number of other papers in the volume are almost strict biostratigraphic studies (for example, Aberhan et al.), again reflecting both the excellent and diverse record of the bivalves, and Newell’s influence on the field. There are several papers that span the broad range of paleon- tological subdisciplines, bringing together numerous ar- eas of thought. One of the finest examples in this volume is the paper by Anneli et al., which combines taphonomic and functional morphological analyses with paleoenviron- mental interpretation to reconstruct bivalve ecology in the Late Paleozoic Paranaiba and Parana Basins of Brazil. While at first glance this volume may appear to be a hotchpotch of interests, brought together only by the common thread of bivalve biology and paleontology, it represents the state of the field today. Studies of bivalves, and studies using bivalves, are of paramount importance in a number of disciplines. Perhaps it is too soon to ex- pect a comprehensive and highly structured volume ded- Page 104 icated to explaining bivalve evolution. But what better tribute could have been assembled for honoring the di- verse and varied interests of a contributor such as Norman Newell? Peter D. Roopnarine Atlas of the Land and Freshwater Molluscs of Britain and Ireland by MICHAEL KERNEY. 1999. Harley Books, Martins, Great Horkesley, Colchester, Essex CO6 4AH, England. 264 pp. ISBN 0-946589-48-8. £25.00. Begun in 1961 (but with intellectual roots that extend back into the prior century), the Conchological Society of Great Britain and Ireland’s project to map the land and freshwater mollusks of the British Isles on a grid of 10- km squares is one of the most ambitious and thorough such mapping schemes ever undertaken. Good coverage of the British Isles has now been achieved, so that the reported ranges are in all probability quite complete, and the differences in apparent regional species diversity re- flect true variation ascribable to climate, topography, ge- ology, or historic happenstance. The dynamic nature of molluscan distribution is thoughtfully discussed in the The Veliger, Vol. 43, No. 1 opening essays, including factors that influence presence and absence, the roles of postglacial climatic history and human activity, and a prospectus for the future. The atlas treats each species individually, giving for each the scientific and vernacular names, common syno- nyms, and size range; an excellent illustration (of the shell for snails and clams, and of the habitus of a crawling individual for slugs); a dot-distribution map based on data in hand as of 1998; and a few paragraphs on habitat, history, and survival status. Fossil occurrences are noted and mapped with special symbols, as are grid records made prior to 1965 only. In some cases, the latter indicate significant recent changes in species’ distribution. Ranges outside the British Isles are summarized. Parallel maps illustrate environmental factors such as January and July mean temperatures, annual rainfall, the distribution of calcareous rocks, and atmospheric sulfur dioxide concentrations. The work concludes with a list of the names of recorders who have participated in the map- ping scheme, and an extensive bibliography with over 200 entries. Not only is this book beautifully produced and data- rich, it serves as a model of what persistence and a ded- icated group of well-organized specialists can accom- plish. We hope, along with the author, that it comes to serve actively as a record against which future change is measured and, perhaps to some extent, controlled. B. Roth Information for Contributors Manuscripts Manuscripts must be typed, one side only, on A4 or equivalent (e.g., 842” X 11”) white paper, and double-spaced throughout, including references, figure legends, footnotes, and tables. All margins should be at least 25 mm wide. Text should be ragged right (i-e., not full justified). Avoid hyphenating words at the right margin. Manuscripts, in- cluding figures, should be submitted in triplicate. The first mention in the text of the scientific name of a species should be accompanied by the taxonomic authority, in- cluding the year, if possible. 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Submitting manuscripts Send manuscripts, proofs, books for review, and corre- spondence on editorial matters to Dr. Barry Roth, Editor, 745 Cole Street, San Francisco, CA 94117, USA. CONTENTS — Continued Distribution of the bonnet limpet, Hipponix conicus (Gastropoda: Hipponicidae), among host species in western Kyushu, Japan KAZUNORI AMAHIRAVAN DW UME @ WAN@ (500) shaven eleire) chal a Manertejaretcy satel eter aye aeee stone Growth and fecundity of Lymnaea elodes (Gastropoda: Lymnaeidae) under laboratory conditions LAUREN) FLORIN, BERNARD FRIED) AND ADITYA-REDDY Yh aiqe a: sleeker ane A new subspecies of the schoolmaster gonate squid, Berryteuthis magister (Cephalopoda: Gonatidae), from the Japan Sea OLEG: Ne KATUGING (a site ote a sth 40) orgies ar oe tole ietae salen dein CUS afer scl Rt Ree NOTES, INFORMATION & NEWS Arboreal Neritidae ROBERT He: GOWIEVAND BARRY. D)s/SMINGHE | 2 oe eee Si eee rere Paleogeographic implications of late Paleocene Onestia onestae (Bivalvia: Cardiidae) in arctic Alaska JAYA. SCHNEIDER AND’ LOUIE: MARINCOVICH, JRE- 3 fan tancle cee san nee BOOKS, PERIODIGALS é¢ PAMPHIEE WS 2) icici. os els raeheeete biens ears t See eee VL 4o\ Vv4yK cae WG MELIGER i A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler, Founding Editor ISSN 0042-3211 } a) ii ' ABRs, Volume 43 April 3, 2000 Number 2 CONTENTS Nocturnalism in Ap/ysia oculifera (Adams & Reeve, 1850): an avoidance behavior minimizing exposure to ultraviolet radiation? | ETAT TRTCANUFIES eet aaa GA sears rts cee ces oe Gy Sk icy GUC RGR aa actae ett oa em a Spiomenia spiculata, gen. et sp. nov. (Aplacophora: Neomeniomorpha) collected from the deep waters of the West European Basin IZ AINIBIEAGAIRIN @ ES KeYaue tants case 2 oe eiterce sane al a Ue Ee emh pe een anal hcl lien ane ik ty ci Mee Sie aie The buccinid gastropod Deussenia from Upper Cretaceous strata of California RICHARDS bs S OUIRESFAN DEO UE HUAU RE: SAUTE rueneinns sins, arsisieet aiiclens chats sich ere ee « 118 Small, high-spired pulmonates from Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with description of a new genus, and with conservation statuses of 15 streptaxid species KENNEDEEC | EMBERTON-AND MIMOMHY AG PEARCE 0.) 5/2. a ene acne es esse ee os 126 Three new Pacific species of Halgerda (Opisthobranchia: Nudibranchia: Doridoidea) Orley CARUSONPAN DNL aa AORBR sey nage etsee nach Gtaierd iG Aiea, soc ce lem asi 8 F siellerele 154 Taxonomic revision of the common Indo-West Pacific nudibranch Phyllidia varicosa Lamarck, 1801 AT EXANDER FARIRNERVAND, MIGHAEL SGHRODIE, «6 dnc sae ecie scl gele false tse. eis 0% 164 Redescription and range extension of Bathydoris aioca Marcus & Marcus, 1962 (Nudibranchia: Gnathodoridoidea) ISN GEIAVANDES MANDI FIAN SI DERUSCH a4 Gee ee oe ce eae ita iee claws «cbse 1772 CONTENTS — Continued The Veliger (ISSN 0042-3211) is published quarterly in January, April, July, and October by the California Malacozoological Society, Inc., % Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. 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Single copies of an issue are US $25.00, postage included. A limited number of back issues are available. Send all business correspondence, including subscription orders, membership applications, pay- ments, and changes of address, to: The Veliger, Dr. Henry Chaney, Secretary, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105, USA. Send manuscripts, proofs, books for review, and correspondence regarding editorial matters to: Dr. Barry Roth, Editor, 745 Cole Street, San Francisco, CA 94117, USA. © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). The Veliger 43(2):105—109 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 Nocturnalism in Aplysia oculifera (Adams & Reeve, 1850): An Avoidance Behavior Minimizing Exposure to Ultraviolet Radiation? ITAI PLAUT! Department of Biology, University of Haifa at Oranim, Tivon, 36006, Israel Abstract. Circadian activity patterns and preferred location were recorded under experimental conditions in two groups of the sea hare Aplysia oculifera, one exposed to direct sunlight (group L) and the other placed in the shade (group S). Observations were made several times during the day and night for 20 days. Activities (resting, feeding, crawling, copulating and spawning) and locations (buried in the sand, under stones, on the substrate and on the algae) were recorded. Data were pooled into four time blocks (midnight to 06:00, 06:01 to 12:00, 12:01 to 18:00, and 18:01 to midnight). Sea hares of group L showed distinct nocturnal activity pattern, feeding at night when most of them were found on the algae, and resting during the day when they were located mostly under the stones. In group S no significant differences were detected between day and night activities or preferred location. The results show that the nocturnal activity pattern in the sea hare A. oculifera is governed by an external factor, probably direct sunlight, rather than an internal cue. It is suggested that A. oculifera is nocturnally active to avoid exposure to direct ultraviolet radiation. INTRODUCTION Aplysia oculifera (Adams & Reeve, 1850) is a widespread sea hare species, distributed in tropical and subtropical regions from the eastern coast of Africa, including the Red Sea, through the Indo-Pacific Ocean to the islands of Hawaii. Aplysia oculifera is a medium-sized sea hare (5-100 g live body weight), which dwells on shallow rocky shores, and feeds exclusively on macroalgae, main- ly the green algae Ulva spp. and Enteromorpha spp (Plaut, 1993; Plaut et al., 1998). Among sea hares of the genus Aplysia, some species are nocturnal and others diurnal (Susswein et al., 1984; Carefoot, 1987, 1989; Carefoot & Taylor, 1988). It is as yet unknown whether the cue for these patterns of activity is endogenous or exogenous, and what advantages sea hares gain by being either nocturnal or diurnal. Carefoot & Taylor (1988) listed several possible rea- sons for nocturnalism in A. dactylomela Rang, 1828 (which ecologically resembles A. oculifera), such as avoiding predators, avoiding diurnal interspecific or con- specific competitors, and intolerance to light, particularly ultraviolet wavelengths. They strongly rejected avoidance of predation and competition as possible reasons because (1) Aplysia has few or no predators (Carefoot, 1987), and instances of predation known for A. dactylomela are iso- lated events of apparent opportunism involving only ju- veniles and eggs being eaten (Willan, 1979). In addition, the defensive repertoire of adult Ap/ysia includes purple ink and opaline gland secretions, unpalatable skin, and ' Phone: +972-4-9531026, fax: +972-4-9832167, e-mail: plaut @research.haifa.ac.il toxic digestive gland; (2) most interspecific competitors seem to show no effect on algae abundance or are noc- turnal as well (e.g., Hobson, 1974); and (3) conspecific competition will not be avoided if the entire population is nocturnal. These same arguments apply also to A. ocu- lifera in the Gulf of Eilat (Plaut, 1993). Carefoot & Taylor (1988), and Carefoot (1989) also rejected the avoidance of ultraviolet radiation as a possible reason because (1) light is unlikely to have operated as a primary force lead- ing to the evolution of nocturnalism; (2) Aplysia makes no obvious attempt to protect its eggs from direct sun- light; and (3) other tropical and subtropical sea hares in- habiting shallow water are diurnal. They suggested that there may be a metabolic advan- tage to splitting the day into two phases of activity (feed- ing and resting) but could not explain the advantage of nocturnalism versus diurnalism or vice versa. Surveys conducted in the Gulf of Eilat (Aqaba) show that A. oculifera is mostly nocturnal (Plaut, 1993; Plaut et al., 1998). Its nighttime activities include feeding, crawling, copulating, and spawning. During the daytime, sea hares mostly rest either under stones or buried in the sand. Contrary to this general observation, in several cas- es, A. oculifera in the Gulf of Eilat was observed feeding, crawling, copulating, and spawning during daytime when algae were very abundant and when it was found at a depth below 2 m (Plaut, 1993). In this study I examined the question of whether the nocturnalism of A. oculifera is a response to external cue, e.g., direct radiation (sunlight), or whether it is governed by internal cues. I followed the locations and activities of two groups of A. oculifera under outdoor experimental Page 106 conditions during daytime and nighttime, one exposed to direct natural sunlight and the other under shade. MATERIALS AnD METHODS Newly recruited (>1 g live body weight at the beginning of the settlement season) sea hares were collected at sev- eral sites in the northern Gulf of Eilat (Aqaba) in Feb- ruary 1990. They were kept at the Interuniversity Institute for Marine Sciences of Eilat in an open circulated sea- water system. Experiments were initiated within 1—2 days after collection. Two groups of 20 individuals of similar size (0.8—1.9 g body live mass) were reared in two un- covered 30 L glass aquaria (SO X 25 X 25 cm length, width, and height, respectively). Both aquaria were placed outdoors. One was in the shade (group S), under a roof and with one of its sides a wall facing west, and the other three sides left open; the other was under direct natural sunlight (group L). Each aquarium was supplied with continuous water flow (2—3 L min~!) to maintain similar temperatures (day water temperature, 26 + 1 °C and night temperature 22 + 1 °C with no significant dif- ferences between the two aquaria). Each aquarium con- tained a layer of 3—4 cm sand from the sea hares’ natural habitat, and two stones, 10—13 cm in diameter, that were placed as shelters for the sea hares. Food (Ulva sp.) was provided in amounts sufficient for ad libitum feeding, but at small rations and in small pieces so as to prevent sea hares from using them as shelters against sunlight. Lo- cation of the sea hares and their activities were recorded randomly four to six times daily (day and night) during 20 days. Four areas were defined in the aquaria for the location of the sea hares: (1) under stone; (2) buried in the sand; (3) on the substrate (including sand, stone, and aquarium walls); and (4) on the algae. Five different ac- tivities were defined (after Carefoot, 1989): (1) resting (inactive); (2) feeding (biting or chewing); (3) crawling (moving around without feeding, spawning or copulat- ing); (4) spawning (laying eggs); and (5) copulating. In several cases, when sea hares were observed conducting more than one type of activity simultaneously, (e.g., feed- ing and copulating, copulating and spawning), all activi- ties were recorded for each individual. The results were treated for each location or activity as percentages of all the sea hares in the aquarium. Although the objective of this study was to detect whether sea hares show different patterns of activities and locations during the day and the night, the results were pooled to four 6 hr time blocks. This was done to refine the detection and to obtain results comparable to those of Carefoot (1989). Times blocks were: (1) from 00:00 to 06:00; (2) 06:00 to 12:00; (3) 12:00 to 18:00; and (4) 18:00 to 24:00. The experiment lasted 20 days. Statistical analyses of the results were made after square root transformation of the percentages, and in- cluded two-way ANOVA between day and night and be- The Veliger, Vol. 43, No. 2 tween treatments and Tukey tests between time blocks and between treatments for each location and for each type of activity. Analyses were performed with Systat 5.04 for Windows (Wilkinson, 1990). Significant differ- ences were declared when P < 0.05. RESULTS In order to compare general patterns of location and ac- tivities between sea hares in the light and those in the shade, data were pooled for each treatment for the whole day (24 hr). This comparison shows that sea hares in group L were significantly more likely to be found buried and under stones and significantly less on the algae. Moreover, they rested more and fed and copulated less than sea hares in group S (Figures 1, 2, P < 0.0001). Dividing the day into four blocks of 6 hours each dem- onstrated more clearly the circadian patterns of sea hares’ location (Figure 1) and activities (Figure 2). Aplysia oculifera were rarely found buried in this ex- periment. However most of the records of buried sea hares were collected in group L during daytime (7.1 + 1.4% and 7.1 + 1.8% from 06:00 to 12:00 hr and 12:00 to 18:00 hr, respectively). At these time blocks they bur- rowed significantly more than during night hours (0% during the night), and significantly more than the sea hares in group S (1.8 + 0.7% between 18:00 to 24:00 h and 0% in the other time blocks). The preferred location of resting sea hares was under the stone; sea hares were found under stones mainly in group L during the day (58.6 + 3.4% from 06:00 to 12: OO hr and 56.4 + 2.3% from 12:00 to 18:00 hr), signifi- cantly more than in the same group at night (less than 5.8%) and significantly more than in group S during all 24 hr (2.1 + 1.4% between 06:00 and 12:00 hr and 0% in the rest of the day). Sea hares were found on the substrate during all times and activities. In group L, 35.0 + 2.0% of the sea hares were on the substrate between 00:00 and 06:00 hr, sig- nificantly more than at all other times and more than group S which spent only about 25% on the substrate. Sea hares were usually located on the algae while feed- ing or copulating. In group S no significant difference between the four time blocks was detected (values ranged between 69.6 and 77.7%). These values were similar to those of group L during the night (61.5 + 1.4 and 71.9 + 2.0% from 18:00 to 24:00 hr and from 00:00 to 06:00 hr, respectively). However, during the day, only 11.8 + 2.6% (06:00 to 12:00 hr) and 17.9 + 1.6% (12:00 to 18: OO hr) of the sea hares in group L were found on the algae, significantly less than the above. Patterns of activities within the groups were somewhat more complex. Sea hares were observed resting mostly in group L during the day (more than 60%), significantly more than in the same group at night (about 3%), and more than in group S (< 5.9% during day and night). I. Plaut, 2000 80 + All day (pooled) 60 4 40 5 4 - es a J 80 00:00-06:00 12:00-18:00 Percentage Ss Oo SS niall (0) ese] 80 18:00-24:00 ” -_._ 12 = oO Meo} oO 2 g 5 & =} nm S oS faa) o o és 2 b=) tx — [=| 3 = e) 5 (oe) Locations Figure | Locations of sea hares Aplysia oculifera from the Gulf of Eilat, Red Sea, under natural photoperiod (group L, clear bars) and in the shade (group S, shaded bars) in the aquaria at four time blocks of the day. Bars indicate mean + SD percentage of sea hares in each group in each location. Feeding showed almost the opposite pattern. In group L only 13.2 + 2.1% (06:00—12:00 hr) and 19.3 + 2.2% (12:00—18:00 hr) were found feeding during the day, sig- nificantly less than during the night (58.8 + 3.1 [18:00— 24:00 hr] and 60.0 + 2.8% [00:00—06:00 hr]). In group S significantly higher percentages of feeding sea hares were recorded throughout the 24 hr than in group L dur- Page 107 All day (pooled) 60 — 40 4 i fe or ee 0+ 80 5 00:00-06:00 ea a ae 20 get Ore 06:00-12:00 Ey ie 5 40 4 20S Aa 0 =e 80 : 12:00-18:00 60 40 4 20 i a al 80 18:00-24:00 ES |p| | pe] | 20 0 | I 2 2 = a 2 B 3 = =] a a4 (Zh eS a 3 1S) } in cine i) Activity Figure 2 Activities of sea hares, Aplysia oculifera from the Gulf of Eilat, Red Sea, under natural photoperiod (group L, clear bars) and in the shade (group S, shaded bars) in the aquaria at four time blocks of the day. Bars indicate mean + SD percentage of sea hares in each group in each activity. ing daytime. Percentages of feeding sea hares were sim- ilar in both groups during the night. From 06:00—12:00 hr only 53.8 + 2.6% of the sea hares in group S were feeding (significantly less than during 18:00—06:00 hr in both groups) and 70.4 + 1.5% between 18:00—24:00 hr, significantly more than all other time blocks in both groups. Crawling activity was sometimes observed coupled with some other activity (feeding, copulating, or spawn- ing). It did not show a distinct nocturnal pattern. In group L, crawling was observed significantly more during the night (16.2 + 1.7% at 18:00—24:00 and 11.9 + 2.1% at 00:00—06:00) than in the daytime (6.8 + 1.4% at 06:00— Page 108 12:00 and 8.2 + 1.3% at 12:00—18:00). In group S crawl- ing was observed most frequently at 12:00—18:00 (18.9 + 2.8%), significantly more than at night (8.8 + 1.7% and 5.0 + 1.0% at 18:00—24:00 hr and 00:00—06:00 hr, respectively). During the 06:00—12:00 time block, sea hares from group S crawled significantly less than in all other time blocks (1.4 + 0.7%). Sea hares in group L showed significant differences between day and night in the frequency of copulation. During the daytime, only 14.3 + 3.8% and 11.4 + 3.0% (at 06:00—12:00 hr and 12:00—18:00 hr, respectively) were copulating, significantly lower than at night (29.9 + 1.9% and 28.8 + 2.7% at 18:00—24:00 hr and 00:00—06: OO hr respectively). In group S copulation activity was similarly high during the night and the first part of the day (35.4 + 2.2%, 42.1 + 4.3% and 23.1 + 3.5% at 00: 00-06:00 hr, 06:00—12:00 hr and 18:00—24:00 hr, respec- tively) and significantly lower in the second part of day- time (18.6 + 2.7% at 12:00—18:00 hr). Unlike all other activities, sea hares in both groups spawned only at the second part of the night (00:00—06: OO hr, 1.5 + 0.7% and 2.3 + 0.8% in group L and group S, respectively) and the first part of the day (06:00—12: OO hr, 3.2 + 0.9% and 4.5 + 0.9% in group L and group S, respectively). Group L continued to spawn in the sec- ond part of the day (12:00—18:00 hr, 1.4 + 0.7%), sig- nificantly less than in the other time blocks. During the 18:00—24:00 hr time block, no spawning was observed. DISCUSSION The main finding of this study is that A. oculifera dem- onstrates a nocturnal activity pattern only when exposed to direct sunlight during the day. It has been clearly shown that when under direct sunlight A. oculifera was quiescent during the day and active at night. When under the shade, exposed to indirect light of natural photoperi- od, where there were, at least, potential light-related cues as to the time of the day, they showed no pattern of noc- turnalism, and were active throughout the whole day and night. Plaut et al. (1996) showed that under shade A. oculifera grew somewhat faster than under direct sunlight. The fact that more than 60% of the A. oculifera individuals under shade were observed feeding throughout the day (Figure 2) makes it doubtful that there is any energetic advantage for A. oculifera being quiescent for more than about 30% of the day, as was suggested by Carefoot & Taylor (1988). Moreover, under direct sunlight, more than 70% of the sea hares were inactive and were located in the shade (under stone or buried). About 20% of the sea hares were on the algae, possibly in partial shade. In addition, A. oculifera under natural photoperiod of direct sunlight grew at arate somewhat slower, although not significantly different than under shade (Plaut et al., 1996). The slower growth rate of sea hares, coupled with greater egg pro- The Veliger, Vol. 43, No. 2 duction under direct sunlight (Plaut et al., 1996) suggests that they may have been feeding less and were under suboptimal conditions of restricted feeding (Plaut et al., 1996). Thus, no energetic advantage of being quiescent for part of the day can be claimed in this case. The fact that sea hares under shade showed neither a nocturnal nor a diurnal pattern of activity suggests that the cue for nocturnalism of sea hares in their natural hab- itat is exogenous. It seems that in this case, as reported previously (Jacklet, 1976; Carefoot & Taylor, 1988; Care- foot, 1989) direct sunlight is the cue for this pattern. Sun- light containing ultraviolet wavelengths has been found to directly harm tropical shallow-water organisms. Dam- age may include decrease in respiration, growth rates, and calcification rates, and death (Shick et al., 1996). Indi- rectly, ultraviolet radiation may damage shallow-water organisms via photochemical reactions that produce re- active oxygen molecules like H,O, (Shick et al., 1996). The natural concentration of stratospheric ozone, gener- ally less near the Equator than at higher latitudes (Cutch- is, 1982), together with the lower solar zenith angle in tropical regions, means that the tropics receive more ul- traviolet radiation. Thus, tropical ecosystems have an evolutionary history of exposure to high fluxes of ultra- violet radiation (Green et al., 1974; Frederick et al., 1989). The transparency of tropical seawater allows pen- etration of ultraviolet radiation in shallow-water habitats (Kirk, 1994). Thus, if ultraviolet radiation may harm sea hares, as it does other organisms in tropical shallow wa- ter, an evolution toward avoidance of being exposed to this radiation is highly probable. Carefoot & Taylor (1988) and Carefoot (1989) consid- ered avoidance of light, particularly ultraviolet wave- lengths, as a possible advantage of nocturnalism in shal- low-water populations of tropical Aplysia. However, they raised three arguments against this hypothesis. First, “light is unlikely to have operated as a primary force leading to the evolution of nocturnalism.’’ Second, A. dactylomela, as A. oculifera (Plaut, 1993) makes no ob- vious attempt to protect its eggs from the sun’s rays. Fi- nally, many other species of tropical and subtropical sea hares are active during the day in shallow water. As for Carefoot & Taylor’s (1988) first argument, it is a general statement, not supported by any explanation and thus it remains unclear. Regarding the argument about unprotected egg masses, A. oculifera, like other sea hares, lays its egg masses dur- ing day and night (Figure 2) wherever it happens to be located while spawning, either exposed to (Carefoot & Taylor, 1988) or hidden from sunlight (Plaut, personal observation). Rawlings (1996) stated that many benthic marine invertebrates, living in tidal and subtidal habitats, including Mollusca, shield their embryos from direct ex- posure to ultraviolet radiation by a capsule wall which effectively filters ultraviolet radiation, as in the caeno- gastropod, Nucella emarginata (Deshayes, 1839). Re- I. Plaut, 2000 Page 109 cently, Carefoot et al. (1998) also suggested possible UV protection in eggs of the sea hare, A. dactylomela. This may be also the case in the egg masses of Aplysia ocu- lifera, indicating evolutionary adaptation of UV avoid- ance. The fact that there are populations of sea hares that do not show a nocturnal pattern of activity should be tested for each population individually. Plaut (1993) reported diurnal activity of A. oculifera in several cases, in all of which sea hares were, at least partially, protected from direct sunlight, either by high amounts of algae, or by being in relatively deep water (> 2 m). The same may apply to other populations of sea hares observed to be active diurnally, thus strengthening the assumption that nocturnalism in sea hares is a form of an opportunistic behavior, directly aimed at avoiding exposure to ultravi- olet radiation. The only activity that showed a constant circadian pat- tern in both groups was spawning, which occurred only between midnight and noon. This pattern may be related to avoidance of egg predators before the egg capsules are fully developed. However, results are insufficient to ex- amine this hypothesis. Acknowledgments. The author wish to thank Mrs. M. Plaut for helping in the observations, Prof. M. E. Spira for helpful dis- cussion, the staff of the Interuniversity Institute of Marine Sci- ences in Eilat, Israel, for hospitality and technical assistance, Dr. A. Genin for radiation data, and Prof. A. Haim for his comments after reading the manuscript. LITERATURE CITED CarEFOOT, T. H. 1987. Aplysia: its biology and ecology. 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Evanston, Illinois. 675 pp. WILLAN, R. C. 1979. The ecology of two New Zealand opistho- branch molluscs. Ph.D. Thesis, University of Auckland, Auckland, New Zealand. 198 pp. The Veliger 43(2):110—117 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 Spiomenia spiculata, Gen. et Sp. Nov. (Aplacophora: Neomeniomorpha) Collected from the Deep Waters of the West European Basin PAMELA ARNOFSKY'! Woods Hole Oceanographic Institution, M.S. #34, Woods Hole, Massachusetts 02543, USA (e-mail: parnofsky @ensr.com) Abstract. Spiomenia spiculata gen. et sp. nov. was collected from deep waters of the West European Basin at depths from 1600 to 4400 meters. Spiomenia is placed in the family Simrothiellidae on the basis of the presence of distichous radular bars, which have many denticles during some point in ontogeny, and paired anteroventral radular pockets. It is separated from other genera in the family by the absence of skeletal spicules, denticulation of the radula, and presence of captate upright spicules. INTRODUCTION Spiomenia gen. nov. is a genus of neomenioid aplacopho- ran mollusk which occurs in the benthos throughout the deep Atlantic. It is placed in the Family Simrothiellidae based on the presence of a radula with distichous bars and paired anteroventral radular pockets. The Family Simrothiellidae at present comprises nine other genera of which four are monotypic (Table 1). Of these, three are described from a single sectioned specimen, and infor- mation concerning the spicules and radulae are not re- ported in detail. There are many incongruent morpholog- ical characters within this family and it is probably not monophyletic. For example, Helicoradomenia Scheltema & Kuzirian, 1991, and Sialoherpia Salvini-Plawen, 1978, possess solid epidermal spicules, but spicules are hollow in other genera. Uncimenia Nierstrasz, 1903, was de- scribed as having both captate spicules similar to those of Spiomenia and barbed spicules similar to neomenioids in the family Pruvotinidae. Uncimenia was also reported to lack a radula. All other genera in the family lack barbed spicules and possess a radula. MATERIALS ann METHODS Specimens (105) were collected from several localities in the West European Basin during the 1976 French spon- sored INCAL/CENTOB cruise (Table 2). Type specimens are deposited in the Muséum National d’ Histoire Natu- relle (MNHM) Paris, France. Dissections Lengths were measured along the midline in lateral view. Width measurements were made with the specimen 'Present address: ENSR Consulting & Engineering, 89 Water Street, Woods Hole, Massachusetts 02543 oriented dorsoventrally. Radulae, epidermal spicules, and copulatory spicules were dissected from three animals; six were sectioned for histology. Radulae were dissected by placing the anterior half of the animal in hypochlorite solution. After the tissue was dissolved, the radula was removed with the remaining anterior spicules and washed in distilled water overnight. Spicules were placed on a glass slide, air dried, and permanently mounted in His- tomount®. Radulae were first observed in a drop of glyc- erine for camera lucida drawing, then washed in distilled water and permanently mounted in CMCP-10® for draw- ing under oil immersion. Copulatory spicules were dis- sociated by placing the posterior half of the animal in hypochlorite solution. After the tissue was dissolved, spicules were removed, washed in distilled water, air dried, and mounted in Histomount® for drawing under the camera lucida. Histology Six specimens, previously fixed in formalin and pre- served in alcohol, were post-fixed overnight in zinc for- malin (Presnell & Schreibman, 1997) and embedded in filtered Paraplast®. Sections were cut at 7 wm and stained with hematoxylin and eosin Y or Mallory’s Trichrome and mounted in Histomount®. Terminology and Abbreviations Terminology and abbreviations in this paper follow those of Scheltema & Kuzirian (1991). New terms not mentioned previously in the literature, some specific to Spiomenia are: the radular buttress (BU) which is the thickened lateral portion of each radular tooth (Figure 1D, E). Double denticle (DD) is the lateralmost denticle on the radular tooth and is composed of two denticles fused at the base (Figure 1D). Captate spicules (CA) are those with small peaks at the apical end (Figure 1F). P. Arnofsky, 2000 Page 111 Table 1 Number of species presently known for each genus currently placed in the family Simrothiellidae. Number of Genus Author, date species Simrothiella Pilsbry, 1898 a Cyclomenia Nierstrasz, 1902 1 Kruppomenia Nierstrasz, 1903a 4* Uncimenia Nierstrasz, 1903b Db Biserramenia Salvini-Plawen, 1968 1 Birasoherpia Salvini-Plawen, 1978 1 Sialoherpia Salvini-Plawen, 1978 1 Helicoradomenia Scheltema & Kuzirian, 1991 3 “New Genus P” Scheltema ahs Spiomenia new herein 4* * Some species descriptions in manuscript. SYSTEMATICS Subclass NEOMENIOMORPHA Pelseneer, 1906 Family SIMROTHIELLIDAE Salvini-Plawen, 1978 Spiomenia Arnofsky, gen. nov. Description: Spiny habitus, typically curved due to pres- ervation, 3.5 mm or less in length; dorsofrontal sensory pit obscured by spicules; proboscis large, protrusible; mouth opening at posterior end of vestibule. Pedal pit large; cuticle thick, thickened laterally and ventrally; spicules hollow, upright, many captate, skeletal spicules absent; radula large, longest denticles on buttress, one to many denticles lateral to the radular buttress; radula with one turn into paired anteroventral radular pockets, first- formed teeth not retained; paired ventral salivary glands large, within muscle tissue; mantle cavity with three pairs of long respiratory papillae, often extending outside man- tle cavity. The two autapomorphic characteristics of the genus are: (1) possession of captate upright spicules and (2) from one to many denticles lateral to the radular but- tress. Distribution: Four species, each from geographically separated basins of the Atlantic Ocean belong to the ge- nus Spiomenia; descriptions of three species remain un- published. Spiomenia spiculata is described from the West European Basin (2000—4000 m). The other species are from the North American Basin (2100 m), from both the Canary and Cape Verde Basins (1600-3100 m), and from the Argentine Basin (1600—5000 m). One specimen collected from the Indian Ocean is thought to be a fifth new species, but more specimens of this form need to be collected. It is likely that Spiomenia species are distrib- uted throughout the deep ocean basins. Etymology: Spio from Greek meaning sea nymph, menia from Greek meaning moon, a typical generic ending for neomeniomorph aplacophorans. Table 2 Locality information for Spiomenia spiculata collected from the West European Basin during the INCAL/CENTOB cruise. Latitude north Longitude west Equipment* Degrees Minutes Degrees CP-01 TR S// Sie 10 CP-02 TR / 58.4 10 CP-05 TR 55 0.4 12 CP-06 TR 55 Pep) 12 CP-07 TR 55 3.4 12 CP-08 TR 50 14.7 13 CP-09 TR 50 15.4 13 DS-02 ES 57 58.8 10 DS-05 ES 56 28.1 11 DS-06 ES 56 26.6 11 DS-07 ES 55 0.7 12 DS-08 ES 55 2.0 12 DS-09** ES 55 Vol! 12 OS-02 ES 50 14.4 13 OS-06 ES 47 27.3 9 WS-01 TR 50 19.4 13 WS-02 TR 50 19.3 12 *TR = Trawl, ES = Epibenthic Sled. ** = Type locality. No. of Minutes Depth (m) Date collected specimens 55 2040 16/07/76 p2 42.8 2091 16/07/76 1 29.4 2884 19/07/76 4 40.3 2888 19/07/76 8 46.2 2895 20/07/76 8 13.5 2644 27/07/76 9 15.8 2659 27/07/76 5) 48.5 2081 16/07/76 10 11.7 2503 18/07/76 12 10.5 2494 18/07/76 4 31 2884 19/07/76 1 34.6 2891 19/07/76 3 52.6 2897 20/07/76 17 10.9 2634 30/07/76 4 36.2 4307 09/08/76 2 8.1 2550 30/07/76 12 55.8 2498 30/07/76 3 Total = 105 Page 112 The Veliger, Vol. 43, No. 2 hoy | aL ih P. Armofsky, 2000 Spiomenia spiculata Arnofsky, sp. nov. (Figures 1—3) Material examined: 105 individuals from the Western European Basin (2040 m—4307 m); See Table 2. Holotype (Figures 1A—C, 2J): 2.3 mm long, anterior width 0.7 mm, midbody width 0.7 mm, posterior width 0.6 mm. INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2897 m. Numbered paratypes: Paratype no. 1: INCAL (CEN- TOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5’W, 2897 m, dis- sected specimen, anterior and posterior spicules and rad- ula. Paratype no. 2: INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2897 m, whole animal. Paratype no. 3: INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2897 m, dissected specimen, ante- rior spicules and copulatory spicules. Paratype no. 4*: INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2897 m, histologic sections (seven slides). Paratype no. 5: INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2987 m, histologic sections (15 slides). Paratype no. 6*: INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2897 m, histologic sections (eight slides). Paratype no. 7: INCAL (CENTOB) Loc. 4, DS-05, 56°28.1'N, 11°11.7'W, 2503 m histologic sections (10 slides). Paratype no. 8*: Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2987 m, histologic sections (five slides). Paratype no. 9*: Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2987 m dissected specimen, radula and anterior spicules. Page 113 Lots sent to MNMH for type series OS-06; 4307 m, DS- 07; 2884 m, DS-08; 2891 m, and type locality DS-09; 2897 m. Lots excluded from type series for further investigation: CP—O1; 2040 m, CP-02; 2091 m, CP-05; 2884 m, CP-06; 2888 m, CP-07; 2895 m, CP-082644 m, CP-09; 2659 m, DS-02; 2081 m, DS-05; 2503 m, DS-06; 2494 m, OS-02; 4829 m, WS-01; 2550 m, WS-02; 2498 m.* Diagnosis: Length to 3.1 mm with a mean size of 1.8 mm (n = 105). Narrowest posteriorly; mean index (height:width) at midbody 1:1. Length of cap of large captate spicules 38—48 w and for small captate spicules 10—30 pm. Radula with single turn into paired anteroven- tral radular pockets; five denticles lateral to the buttress, lateralmost being a double denticle; formula 22—25 x 1: 1, 20-23 denticles per tooth (Figures 1D, E, 2J). Two types of copulatory spicules in paired groups protruding through the mantle cavity opening (Figure 3D). External anatomy and hard parts: Body curved, pos- terior end tapering slightly, usually widest at midsection (Figure 1A—C). Dorsofrontal sensory pit, dorsoterminal sense organ and mantle cavity opening obstructed by spicules, difficult to discern. Longest epidermal spicules posterior, near mantle cavity opening. Spicules of nine types (Figure 1F): Type 1, large, captate, 350-450 p.m in length, up to 10 pm in width, hollow to cap; recurved 50 ym from base. Type 2, smaller, captate, 190-240 wm in length, 6 1m in width, hollow to cap. Type 3, large, long, slightly curved, obliquely truncate at apical end, 325—800 wm in length, up to 15 wm in width; recurved 35 pm * Numbered paratypes dissected or histologically sectioned ex- cluded from type series: Paratypes nos. 4, 6, 8, and 9 INCAL (CENTOB) Loc. 4, DS-09, 55°7.7'N, 12°52.5'W, 2897 m. Figure | Spiomenia spiculata Arnofsky, gen. et sp. nov. A—C: Holotype. A. Entire, showing orientation of spicules. B. Anterior end, frontal view showing opening of mouth. C. Posterior end, ventral view, oval-shaped mantle cavity opening obscured by spicules. D—E. Paratype no. 1. Two adjacent teeth of radula, lateral to right. E. Single radular tooth, view from beneath radular membrane showing thickened lateral buttress. EK Epidermal spicules from paratypes nos. 1, 2, and 3: Types 1—6 found both anteriorly and posteriorly, largest of type 3 found near mantle cavity opening, types 7—9 found posteriorly, and types 10, 11, and 12 found ventrally beside pedal groove. G, H. Schematic diagram from histologic sections of two individuals: G. Anterior end to midsection (paratype no. 5). H. Midsection to posterior end (paratype no. 7): transverse sections 1—4 are keyed to sections A—D in Figure 2; transverse sections 5-8 are keyed to sections A—D in Figure 3. I, K. Accessory copulatory spicules, paratype no. 1. I-L Paratype no. 3. I. In situ showing arrangement accessory spicules around mantle cavity opening. Stippling indicates abrasion. J and L. Copulatory spicules. J. Type 1 with sharp bend, remaining hollow portion broken during dissection. K. Accessory copulatory spicule. L. Type 3 found with type 1 spicule in copulatory spicule pocket. Key: BU, buttress; CA, captate spicule; CG, cerebral ganglion; CS, copulatory spicule pocket; DC, dorsal cecum; DD, double denticle; DSO; dorsoterminal sense organ; DSP, dorsofrontal sensory pit; E, eggs; Gd,, upper gametoduct; Gd,, lower ga- metoduct; H, heart; MC, mantle cavity, MG, midgut; MO, mouth; O, oocyte; P, pedal pit; PC, pericardial cavity; PG, pedal gland; PH, pharynx; RB, radula bolster; RP, respiratory papillae; SR, seminal receptacle; VE, vestibule; VS, ventral sinus; VSG, ventral salivary gland. Scale bars: A, B, C = 500 pm; D, E = 10 wm; F top scale bar for spicules 1-6 = 50 pm, bottom scale bar for spicules 7-12 = 25 ym. I-L scale bar = 30 pm. Page 114 The Veliger, Vol. 43, No. 2 2 , ea Ty 4 : fee a) Figure 2 Spiomenia spiculata Arnofsky, gen. et sp. nov. A-E and G-H Paratype no. 5. A—D. Transverse histologic sections 1 to 4 respectively of Figure 1G. A. Section 1 through cerebral ganglion, vestibulum, and pedal gland. B. Section P. Arnofsky, 2000 from base; apical end solid at tip for 20 wm. Type 4, slightly curved, 275-310 wm long, 10 pm in width, bend at 25 wm from base; apical end solid at tip for 25 pm. Type 5, spicule length 75-175 pm in length; widest 20 ym from base, 10—15 wm wide; apical end solid at tip for 10 pm. Type 6, “‘S’’-shaped spicule 220—240 ym in length, 15 wm in width, rounded, solid apical end 25 wm long, recurved at 75 wm from base. Type 7, recurved, apical end pointed, 54 um in length, 6 wm in width. Type 8, small “S”’-shaped spicule, pointed at apical end, 40— 50 pm long, 5 pm in width. Type 9, sword-shaped, solid, found near pedal groove, widest 12 4m from base, 55— 65 pm in length, 6 wm in width. Pedal groove spicules solid with three types. Type 10, oval, with small thick- ened portion at base, length 30—40 pm, 12 wm in width. Type 11, triangular, solid, 35—45 wm long, 9 wm in width. Type 12, solid, curved with thin basal end, ranging from 50-60 pm in length, up to 8 pm in width. Copulatory spicules of two types: type 1 bent and sharply pointed, 100 ym from bend, base hollow (Figure 1J); type 2 dis- tally wavy, hollow, 280 pm in length (Figure 1L). Several accessory copulatory spicules grouped near mantle cavity opening (Figure 11); one type, long, hollow, slightly curved, length up to 250 pm (Figure 1K). Radula morphology: Radulae from eight animals ex- amined (Figures 1D, E, 2J). Radula distichous, formula 22-25 X 1:1 with 20-23 denticles per tooth. Most re- cently formed teeth 85—90 ym in greatest dimension. Five denticles lateral to buttress, double denticle 7 ym; single lateralmost denticles 7.5—-8.5 m. Three large denticles on buttress 13.5—16.0 ym in length. Smaller denticles me- dial to buttress, 15—16 in number, 0.4—8.5 ym in length on younger teeth, width of radular bar at widest portion 14 wm, rest of bar 10.6 pm. Internal anatomy: Cuticle thickest at anterior end, thick- ened laterally and ventrally throughout, thinnest dorsally at midsection measuring 10 pm thick, ventral thickness measuring 40 pm. Epidermis 10—20 pm thick, with more than one type of secretory cell. Body-wall musculature well developed with longitudinal, transverse, and diago- nal muscles (Figure 2D). Pedal pit lined by large secre- tory cells. Foot small, ventral longitudinal muscles pro- nounced (Figure 3A, C). Vestibular papillae absent. Oral Page 115 cavity with narrow mouth opening to vestibulum (Figure 2B). Dorsal salivary gland large and paired (Figure 2C, D). Anteroventral radular pocket as paired pouches. Rad- ula bolsters large, radular muscles well developed (Figure 2C-E) and paired ventral salivary glands within radular musculature (Figure 2D, E). Short esophagus and single, large dorsal midgut cecum (Figure 2C—E). Oocytes nu- cleated in gonad anterior to pericardial cavity (Figure 2F). Gonopericardial ducts paired until just before opening into pericardial cavity. Heart free within pericardium, pericardial cavity large, filled with yolky ova, which lack a nuclear membrane or nucleolus (Figures 1H, 3A). Opening of upper pericardium into upper gametoduct not determined. Seminal receptacles bilobed, paired, opening into upper gametoducts through a narrow duct (Figure 3A). Paired upper gametoducts convoluted. Lower ga- metoduct single, with a lower lobe into which empty up- per gametoducts (Figure 3B). Seminal vesicles are absent. Gametopore single, opening into mantle cavity below rec- tum. Upper wall of anterior mantle cavity filled by large, vacuolated epithelial cells (Figure 3B). Dorsoterminal sense organ small, seen only in sectioned material. Remarks: This species has unusually large, paired, dorsal salivary glands that empty into the esophagus where the it joins with the radular sac. The ventral salivary glands are also unusual in that they are embedded within the thick layer of muscle tissue that surrounds the radula. Nematocycts were not found in the gut of the six sec- tioned animals (Figure 2G, H). However, diatoms and un- identified spicules, which resemble those of sponges, were found in the vestibule area and within the digestive system, suggesting that this species may not depend on cnidarians as a food source. Retention of ova in the pericardial cavity is unusual among the Aplacophora and has not been noted in the Simrothiellidae. Oocytes have a nuclear membrane and nucleolus in the gonad; however, once ova are within the pericardial cavity, the nuclear membrane has broken down. The seminal receptacles empty into the upper ga- metoducts, so it may be possible that sperm travel through the upper portion of the upper gametoducts into the pericardial cavity and there fertilize the eggs. It is also possible that breakdown of the nuclear membrane pre- 2 through oral cavity, mouth opening, and vestibulum. C. Section 3 through thick radular muscles, radula, dorsal salivary gland, and dorsal cecum. D. Section 4 through paired ventral salivary glands embedded in the thick circular muscle layers of radula; dorsal salivary gland, dorsal cecum, longitudinal and circular muscles are also indicated. E. A more posterior section through the ventral salivary glands and radula showing the radular membrane. F Paratype no. 7. Section through gonad with yolky oocytes. G, H. Contents of midgut, diatoms indicated by arrows. I. Paratype no. 1. Radula. J. Holotype. Key: BU, buttress; CG, cerebral ganglion; CMU, circular muscle; CU, cuticle; DD, double denticle; DC, dorsal cecum; DSG, dorsal salivary gland; G, gonopericardial duct; LMU, lon- gitudinal muscle; MO, mouth; NU, nucleolus; O, nucleus; OC, oral cavity; PG, pedal gland; R, radula; RM, radular muscle; VE, vestibulum; VSG, ventral salivary gland; Y, yolk. Scale bars: A, C, D, and E = 40 um; B, EF G, and H, = 10 pm; I = 40 pm; J = 500 pm. Page 116 The Veliger, Vol. 43, No. 2 Figure 3 Spiomenia spiculata Arnofsky, gen et. sp. nov. A—D Paratype no. 7. Transverse histologic sections through posterior end; A—D represent sections 5—8 respectively of Figure 1H. A. Section 5 through seminal receptacle, lower ga- metoduct, upper gametoduct, heart, and pericardial cavity with large ova perhaps being brooded. B. Section 6 through rectum, mantle cavity, and connection of upper gametoduct with lower portion of lower gametoduct. C. Section 7 through mantle cavity with paired respiratory papillae, arrows, lumina of accessory copulatory spicules. D. Section 8 through posteriormost portion of mantle cavity illustrating the unusual size of the respiratory papillae. Scale bars: A-C = 40 pm, D = 10 pm. Key: AS, accessory copulatory spicules; CS, copulatory spicule pocket; CU, cuticle; F foot; H, heart; GD,, lower gametoduct; GD,, upper gametoduct; MC, mantle cavity; OV, ovum; PC, pericardial cavity; RP, respiratory papillae; SR, seminal receptacle. P. Arnofsky, 2000 Page 117 cedes fertilization. The origin of the upper gametoduct was not determined, as the population was comprised mostly of juveniles and it was difficult to find fully de- veloped adults. However, it is certain that the pericardial cavity is closed at the posterior end and therefore the upper gametoduct must originate toward the anterior por- tion of the pericardial cavity (Figure 1H). Eggs from the pericardial cavity in living Epimenia australis do not de- velop and presumably have not been fertilized. Those be- ing brooded in the mantle cavity, where they are held together by a sticky substance, develop into larvae (Scheltema & Jebb, 1994). It may be possible that the epithelial cells within the mantle cavity of Spiomenia also produce a mucus to bind eggs. Relationships: Spiomenia spiculata radulae were com- pared with the radulae of the nine other genera that have previously been placed in the family Simrothiellidae. Iso- lated radulae from Helicoradomenia, Simrothiella, Krup- pomenia, “new genus P”’ (Scheltema, in manuscript), and Unicimenia were examined, but only illustrations of the radulae drawn from histologic sections are available for Sialoherpia, Birasoherpia, Biserramenia, and Cyclomen- ia. All genera except Uncimenia have distichous bars and paired anteroventral radular pockets. The radulae of gen- era Simrothiella, “‘new genus P,”’ and Spiomenia all pos- sess a radular buttress. Kruppomenia also possess a den- ticulate, thickened, lateral portion to the radula. The external morphology of Spiomenia most closely resembles that of “new genus P” (as Simrothiella in Sal- vini-Plawen, 1978). However, “‘new genus P”’ is about as wide as it is long, whereas Spiomenia is longer than wide. Both have denticulate, large radular buttresses and hollow, straight, upright spicules. In Spiomenia, there are several denticles lateral to the buttress; these denticles are absent for “new genus P.” The radula of “‘new genus P”’ is longer and has several turns into the anteroventral rad- ular pockets, whereas radulae of Spiomenia turn only once into the anteroventral radular pocket. Salvini-Plawen (1978) placed the family Simrothiellidae in the order Cavibelonia based on possession of hollow spicules. Scheltema & Kuzirian (1991) determined that the families of Cavibelonia vary in respect to type of radula and ven- tral salivary glands and in presence or absence of skeletal spicules. They also noted that the morphologies of these structures are not unique to the Cavibelonia but are found in other orders as well, and suggested that the order Cav- ibelonia may not be monophyletic. Helicoradomenia has solid upright spicules and has a very different radular morphology from the other genera placed in this family, and it is likely that Helicoradomenia does not belong in Simrothiellidae. Uncimenia, which was figured by Nier- strasz (1903b) with captate spicules similar to those of Spiomenia, also possesses hooked spicules similar to gen- era that have been placed in the family Pruvotinidae. His- tological sections of Uncimenia neopolitana collected near the type locality possess a tiny radula similar to the distichous hooks of the Pruvotinidae (Arnofsky, in man- uscript). In conclusion, it is likely that neither the Order Cavibelonia nor the family Simrothiellidae are monophy- letic. Acknowledgments. Specimens were kindly provided by the Centre National de Tri d’Océanographie Biologique. I thank Dr. Amélie Scheltema for carefully reviewing this manuscript and for her help in guiding me through the process of becoming knowledgeable in aplacophoran systematics and taxonomy. Spe- cial thanks also to Dr. Ernest Ruber, Dr. James A. Blake, and Ethel LeFave for their encouragement and support. This work was funded by NSF-DEB, PEET 95-21930. WHOI contribution number 9747. LITERATURE CITED NIERSTRASZ, H. F 1902. The Solenogastres of the Siboga Expe- dition. Siboga-Expeditie 47. 46 pp. NIERSTRASZ, H. E 1903a. Kruppomenia minima n.g. n.sp. Mitt- heilungen aus der zoologischen Station der Neapel 16:109— 278. NIERSTRASZ, H. F 1903b. Neue Solenogastren. Zoologische Jahr- bucher, Abteilung fiir Anatomie und Ontogenie der Thiere 18:359-386. PELSENEER, P. 1906. A Treatise on Zoology. E. R. Lankester (ed.), Part 5, Mollusca. Black Press: London. 355 pp. PRESNELL, J. K. & M. P. SCHREIBMAN. 1997. Humason’s Animal Tissue Techniques. 5th ed. Johns Hopkins University Press: Baltimore. xix + 572 pp. Pitssry, H. A. 1898. Order Aplacophora v. Ihering. Pp. 281—310 in Tryon’s Manual of Conchology 17. SALVINI-PLAWEN, L. v. 1968. Neue Formen in marinen Mesop- samon: Kamptozoa und Aculifera (nebst der fiir Adria neuen Sandfauna). Annales der naturhistorische Museum Wien 72: 231-272. SALVINI-PLAWEN, L. v. 1978. Antarktische und subantarktische Solenogastres (eine Monographie: 1898-1984). Zoologica 44:1-315. SCHELTEMA, A. H. & M. Jess. 1994. Natural history of a soleno- gaster mollusc from Papua New Guinea, Epimenia australis (Thiele) (Aplacophora: Neomeniomorpha). Journal of Nat- ural History 28:1297—1318. SCHELTEMA, A. H. & A. M. KuzirIAn. 1991. Helicoradomenia juani gen. et. sp. nov., a Pacific hydrothermal vent Apla- cophora (Mollusca: Neomeniomorpha). The Veliger 34(2): 195-203. The Veliger 43(2):118-125 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 The Buccinid Gastropod Deussenia From Upper Cretaceous Strata of California RICHARD L. SQUIRES Department of Geological Sciences, California State University, Northridge 91330-8266, USA AND LOUELLA R. SAUL Invertebrate Paleontology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, USA Abstract. Rare specimens of three new species of the Late Cretaceous buccinid gastropod Deussenia Stephenson, 1941, are reported from California. Deussenia sierrana sp. nov. is from lower Campanian strata in the Chico Formation in the Pentz area, Butte County, northern California. Deussenia californiana sp. nov. is from upper middle to lower upper Campanian strata in the Tuna Canyon Formation in the Garapito Creek area, eastern Santa Monica Mounains, Los Angeles County, southern California. Deussenia pacificana sp. nov. is from uppermost Maastrichtian or possibly lowermost Paleocene strata in the Dip Creek area, northern San Luis Obispo County, central California. These three new species are the only known occurrences of this genus from the Pacific coast of North America. Deussenia has been reported before only from upper Santonian to lower Campanian strata at the mouth of the Mzamba River (Pondoland, Transkei) in South Africa and from Campanian to Maastrichtian strata in Texas and the Gulf Coast of the United States. INTRODUCTION Late Cretaceous buccinid gastropods are relatively un- common on the Pacific coast of North America. Recent inspection of previously collected material resulted in the detection of three new species of genus Deussenia from widely separated locales in California (Figures 1, 2). It is the purpose of this paper to describe and name these spe- cies. They significantly extend the biogeographic range of Deussenia, which was previously known only from Upper Cretaceous rocks in South Africa and southeastern United States. Abbreviations used are: CASG, California Academy of Sciences, Geology Section, San Francisco; CSUC, De- partment of Geology & Physical Science, California State University, Chico; CSUN, Department of Geological Sci- ences, California State University, Northridge; LACMIP, Natural History Museum of Los Angeles County, Inver- tebrate Paleontology Section. SYSTEMATIC PALEONTOLOGY Superorder CAENOGASTROPODA Cox, 1959 Order NEOGASTROPODA Thiele, 1929 Family BUCCINIDAE Rafinesque, 1815 Subfamily MELONGENINAE Gill, 1871 Discussion: Stephenson (1941) questionably assigned his genus Deussenia to family Buccinidae, but he gave no discussion as to why he chose this family. Sohl (1964) assigned Deussenia to family Melongenidae, and he also gave no discussion for the basis of this assignment. Mel- ongenids have pyriform to fusiform shells, usually shoul- dered whorls, a long anterior canal, and a smooth colu- mella (Wenz, 1943; Davies & Eames, 1971; Rosenberg, 1992). These morphologic features are present on all spe- cies of Deussenia Stephenson, 1941, including the new species described here. Ponder & Warén (1988) regarded melongenids to be a subfamily of Buccinidae. Akers & Akers (1997) did likewise and, furthermore, placed genus Deussenia in subfamily Melongeninae. Genus Deussenia Stephenson, 1941 Type species: Deussenia cibolensis Stephenson, 1941, by original designation; Upper Cretaceous (upper Maastrich- tian) Kemp Formation of the Navarro Group, eastern Tex- as. Discussion: Deussenia resembles the Upper Cretaceous bucciniform Aliofusus Stephenson, 1941, but Deussenia differs from Aliofusus by having a stronger subsutural collar, less inclined growth lines on the collar, and straighter axial ribs. Aliofusus has axial ribs that are curved and follow the outline of the outer lip. Deussenia superficially resembles the Upper Creta- ceous volutid genus Volutomorpha Gabb, 1877, but Deussenia lacks volutid characteristics in that it has no R. L. Squires & L. R. Saul, 2000 Figure 1 Index map to type locality areas of the three new species of Deussenia. fold(s) on the columella, no posterior notch, and no sig- moidal deflection of growth lines near the suture. Fur- thermore, Deussenia differs from Volutomorpha by hav- ing a higher spire, a shorter body whorl, a twisted anterior end, and more dense ornamentation. Prior to this present study, only 10 other species have been placed in genus Deussenia, and Sohl (1964:200) listed them. All are of Late Cretaceous age. Four of the species are from upper Maastrichtian strata in eastern Texas, but Sohl (1964) believed that some of these names might be synonyms because they are (1) distinguished on minor differences in shape and ornament, (2) all are from the same stratigraphic horizon in a limited geographic area, and (3) the number of available specimens of these four species are so few that it is not possible to determine whether or not the minor differences are significant. Four of the other known species of Deussenia are found in upper Campanian to Maastrichtian or upper Maastrichtian strata in Texas, Mississippi, and Tennessee, and are found, to a lesser degree, in similar age strata in Alabama and Georgia (Stephenson, 1941; Sohl, 1964; Akers & Akers, 1997). The other two known species of Deussenia are from upper Santonian to lower Campanian strata at the mouth of the Mzamba (= Umzamba) River, Page 119 Age Sub- Deussenia (Ma) Stage stage n. spp. Ranges D. californiana D. sierrana sarowan [ooeer] Figure 2 Age and stratigraphic positions of the three new species of Deus- senia. Geochronologic time scale from Gradstein et al. (1995). Pondoland, Transkei, South Africa (Sohl, 1964). These latter two species were questionably assigned by Sohl (1964) to genus Deussenia. Deussenia sierrana Squires & Saul, sp. nov. (Figures 3—5) Diagnosis: A small-shelled species of Deussenia with a prominent sutural cord, subtabulate body whorl shoulder, and nearly obsolete body whorl sculpture. Description: Shell small, 25 mm high (incomplete); fu- siform; spire of medium height, about two-fifths of total height of shell; spiral angle about 45°. Protoconch not preserved. Whorls 6 (estimated). Upper spire whorls low- ly convex, smooth. Ramp concave, constricted, and well differentiated starting on more mature half of ante-pen- ultimate whorl and continuing onto penultimate and body whorls; ramp widest (about 0.75 mm) on body whorl. Ramp always bordered posteriorly by a prominent sutural cord and always bordered anteriorly by a subtabulate shoulder. Prominent sutural cord somewhat undulatory on body whorl. On penultimate whorl, subtabulate shoulder with low axial ribs. Body whorl sculpture mostly obso- lete, with faint spiral ribbing near neck area on ventral face. Growth lines prosocline on ramp; sharply flexed (sinused) and opisthocine on subtabulate shoulder; broad- ly prosocline on most of body whorl; nearly straight on whorl base; and strongest near outer lip where growth rugae develop. Aperture elongate, anterior end (incom- plete) slightly twisted to left. Columella smooth, with a The Veliger, Vol. 43, No. 2 Page 120 R. L. Squires & L. R. Saul, 2000 light callus. Outer lip sinuous, notched slightly opposite subtabulate shoulder. Dimensions of holotype: Height 25 mm (incomplete at both extremities), width 11.2 mm. Holotype: LACMIP 12717. Type locality: CSUC loc. PN32, latitude 39°39’08’N, longitude 121°35’50’W. Distribution: Chico Formation, informal Pentz Road member of Russell et al. (1986), Pentz area, Butte Coun- ty, northern California. Geologic age: Late Cretaceous (early Campanian). Discussion: Only three specimens were found. Two are from CSUC locs. PN31 and PN32, and the third is from LACMIP loc. 10833. Except for the holotype, the speci- mens are internal molds. It is likely that the holotype has been slightly worn or weathered, but the shell does not appear to ever have been strongly sculptured. Specimens of Duessenia sierrana are known only from the Chico Formation in the informal Pentz Road member. Russell et al. (1986) inferred that the member was de- posited under estuarine conditions and contains mixed shallow-marine and brackish-marine faunal assemblages. Many faunal elements of the Pentz assemblages, however, suggest normal-marine conditions (Squires & Saul, 1997; Haggart et al., 1997), and the localities yielding D. sier- rana were considered by Watkins & Gohre (unpublished MS) to represent a fully marine, shoreface deposit. In the Pentz area, the Chico Formation probably does not exceed 150 m (450 ft.) in thickness (Russell et al., 1986), and based on the presence of the ammonites Bac- ulites chicoensis (Trask, 1856) and Submortoniceras chi- coense (Trask, 1856), the rocks are early Campanian in age (Matsumoto, 1960; Russell et al., 1986). The early Campanian magnetic anomaly 33R is included within the ranges of both of these ammonites (Ward et al., 1983). The gastropod Anchura callosa Whiteaves, 1903, is also present in these rocks, and suggests an early Campanian age (Elder & Saul, 1996). Deussenia sierrana somewhat resembles the so-called Cryptorhytis pseudorigida Rennie (1930:227-228; Page 121 Woods, 1906:321—322, pl. 39, figs. 2a—c; pl. 40, fig. 1) from the Upper Cretaceous Mzamba Formation in South Africa. The genus Cryptorhytis Meek, 1876, is not a buc- cinid because it has one weak fold on the inner lip and a weaker fold in the interior of the aperture, and Wenz (1943) believed it to be a volute. Sohl (1964) was the first to suggest that “C.”’ pseudorigida might belong to Deussenia. We also consider Rennie’s species to be a Deussenia because it has a fusiform shape, a long anterior canal, a smooth columella, and a concave ramp with a cord bordering the suture; all of which characterize Deus- senia. The new species differs from D. pseudorigida by being narrower just anterior to the ramp, by having only a hint of spiral sculpture rather than strong spiral sculp- ture, and by having no axial nodes on the shoulder of the body whorl. Rennie (1930) provided the name Cryptorhytis pseu- dorigida for specimens misidentified by Woods (1906) as the closely allied, so-called ‘‘Cryptorhytis”’ rigida (Baily, 1855:459, pl. 12, fig. 14; Rennie, 1930:225—227, pl. 27, figs. 9-12; [non Woods, 1906:321, pl. 39, fig. 2a—c, pl. 40, fig. 1 = “C.”’ pseudorigida Rennie]). ‘‘Cryptorhytis”’ rigida differs from “‘C.”’ pseudorigida by having a higher spire, a much more tabulate body whorl shoulder, and narrower and more numerous axial ribs. Based on the very close morphological similarities between these two species, we also consider “‘C.”’ rigida to be a Deussenia. Both species are based on specimens collected from the type section of the Mzamba Formation at the mouth of the Mzamba River. Rennie (1930) reported “C.”’ pseu- dorigida from Beds 3 and 16, and Greyling (1992) re- corded “‘C.”’ rigida from Beds 9, 11, and 16. Klinger & Kennedy (1980), on the basis of the ammonite fauna, placed the Santonian-Campanian boundary at the base of Bed 8, but Greyling (1992) placed it at the top of Bed 12. In spite of this debate, both species are of late San- tonian to early Campanian age and are about the same age as D. sierrana. Etymology: The new species is named for the Sierra Ne- vada range, which rises immediately east of Pentz, north- ern California. Explanation of Figures 3 to 12 All specimens coated with ammonium chloride. Figures 3-5. Deussenia sierrana Squires & Saul, sp. nov., ho- lotype LACMIP 12717, CSUC locality PN32, Pentz area, height 25 mm, X2.6. Figure 3. Apertural view. Figure 4. Abapertural view. Figure 5. Right-lateral view showing outline of outer lip. Figures 6-9. Deussenia californiana Squires & Saul, sp. nov., CSUN loc. 153, Garapito Creek area. Figures 6—8. Holotype LACMIP 12718, height 11 cm, <1. Figure 6. Apertural view. Figure 7. Abapertural view. Figure 8. Apertural view with a por- tion of body whorl removed to better show the columella. Figure 9. Paratype LACMIP 12719, apertural view, height 10.75 cm, x0.8. Figures 10-12. Deussenia pacificana Squires & Saul, sp. nov., holotype CASG 61598.01, CASG loc. 61598, Dip Creek area, 1.9. Figure 10. Apertural view. Figure 11. Abapertural view. Figure 12. Right-lateral view. The Veliger, Vol. 43, No. 2 Deussenia californiana Squires & Saul, sp. nov. (Figures 6—9) Diagnosis: A large-shelled species of Deussenia with a rounded body whorl shoulder bearing about 20 low and narrow axial ribs, and with numerous spiral ribs over the entire body whorl. Description: Shell large, up to 13 cm high (estimated); fusiform; spire of medium height, about two-fifths of total height of shell; spiral angle about 50°. Protoconch not preserved. Whorls 6 1/2 (estimated). Spire whorls tabu- late, steep-sided with axial ribs stronger than spiral ribs. Posterior part of body whorl constricted to a moderately broad and lowly concave ramp, with spiral ribs weaker on ramp than on area of greatest inflation of whorl. Body whorl elongate, with greatest inflation from rounded shoulder to medial part of whorl. Body whorl sculptured by about 20, narrow and widely spaced axial ribs, becom- ing obsolete toward medial part of whorl, and numerous and closely spaced spiral ribs over entire body whorl, persisting onto the ventral surface of the neck. Axial ribs more prominent than spiral ribs on posterior half of body whorl. Aperture elongate-lenticular, anterior end twisted to left. Columella smooth. Dimensions of holotype: Height 11 cm (tip of spire and extreme anterior end both missing); width 5.8 cm. Holotype: LACMIP 12718. Type locality: CSUN loc. 153, latitude 34°07'N, longi- tude 118°34’W. Paratype: LACMIP 12719. Distribution: Tuna Canyon Formation, south fork of Gar- apito Creek, eastern Santa Monica Mountains, Los An- geles County, southern California. Geologic age: Late Cretaceous (late middle to early late Campanian) = Metaplacenticeras pacificum ammonite zone. Discussion: Only two specimens have been found, and both are internal molds. The paratype is larger (estimated total height 13 cm, width 6.5 cm), but most of its spire is missing. The paratype shows spiral ribbing on the neck of the body whorl (Figure 9), whereas on the holotype the spiral ribbing in this area is not preserved. Deussenia californiana is similar to Deussenia cibolo- ensis Stephenson (1941:332—333, pl. 64, figs. 13, 14; Ak- ers & Akers, 1997:figs. 183-184) from the Upper Cre- taceous Kemp Clay [also referred to as the Kemp For- mation] in eastern Texas. Modern workers (e.g., Sohl, 1964:fig. 12; Elder, 1996) correlated the Kemp Clay to the upper Maastrichtian Stage. The new species differs from D. ciboloensis by having a much less tabulate body whorl shoulder, no tubercules on the body whorl shoulder, and narrower spiral ribs on the body whorl. Deussenia californiana is also similar to Deussenia pseudorigida (Rennie, 1930:227—228; Woods, 1906:321— 322, pl. 39, figs. 2a—2c; pl. 40, fig. 1) from upper San- tonian to lower Campanian rocks in the Mzamba For- mation in South Africa (See ‘“‘Discussion”’ under D. sier- rana sp. nov.). The new species differs from D. pseu- dorigida by being narrower just anterior to the concave ramp, by having narrower and more numerous narrower axial ribs, and by having weaker spiral ribs on the anterior half of the body whorl. The type locality of Deussenia californiana in the Gar- apito Creek area in the eastern Santa Monica Mountains, Los Angeles County, southern California, plots in carto- graphic unit ““Kss”” (unnamed Upper Cretaceous strata) of Dibblee’s (1992) map, which is the most recently pub- lished geologic map of the region. Rocks belonging to unit ““Kss”’ in the Temescal Canyon-Santa Ynez Canyon just southeast of Garapito Creek were assigned by Col- burn (1996) to the Upper Cretaceous Tuna Canyon For- mation of Yerkes & Campbell (1979). This formation was deposited by turbidity currents on submarine fans (Yerkes & Campbell, 1979, 1980; Dibblee, 1992), and unit ““Kss”’ represents a dominantly sandy facies. Unit “‘Kss”’ in the region of Garapito Creek corresponds to ““member D” mentioned by Popenoe (1973) and to the so-called ‘“‘upper Chico’? Formation utilized by Carey & Colburn (1978). Popenoe (1973:26—27) reported that “‘member D”’ rocks were probably deposited as turbidities and that the mol- lusk fossils are shallow-marine forms that might have been transported from their regular habitat into somewhat deeper water. Carey & Colburn (1978) reported that these same rocks represent middle-fan channelized turbidites containing lenses of concentrated shallow-marine mollus- can shells that appear to have been transported. The paleoenvironment of the Tuna Canyon Formation closely resembles that of the Chatsworth Formation of Colburn et al. (1981) in the Simi Hills just to the north of the Santa Monica Mountains. According to Dibblee (1992), the Tuna Canyon Formation is probably equiva- lent to the Chatsworth Formation in the Simi Hills. Based on the presence of the ammonite Metaplacenti- ceras pacificum sensu stricto (Smith, 1900), Popenoe (1973) assigned a late Campanian age to the rocks at CSUN loc. 153. Subsequent detailed collecting at this type locality of D. californiana yielded this ammonite, as well as the gastropods Anchura phaba Elder & Saul, 1996, Volutoderma magna Packard, 1922, and Zinsitys kingi (Gabb, 1864); the bivalves Crassatella elongata Anderson, 1958, Indogrammatodon sp., Cucullaea sp., Pinna sp., (closed valved), Pterotrigonia evansana (Meek, 1858), Inoceramus sp., Clisocolus dubius (Gabb, 1864), an isognomid, and a venerid. The presence of Me- taplacenticeras pacificum sensu stricto is very age diag- nostic because this species constitutes the Metaplacenti- ceras pacificum ammonite zone (after Matsomoto, 1960), which is of middle to early late Campanian age (Elder & R. L. Squires & L. R. Saul, 2000 Page 123 Saul, 1996). The geologic ranges of both Zinsitys kingii and Anchura phaba are correlative to the M. pacificum zone (Saul, 1988; Elder & Saul, 1996). At LACMIP loc. 27002, which is in the general area of CSUN loc. 153, the bivalves Glycymeris (Glycymerita) veatchii (Gabb, 1864), Pterotrigonia evansana (Meek, 1858), Cymbophora triangulata (Waring, 1917), Calva sp., and Yaadia sp. were also found. The exact location of LACMIP loc. 27002 is not known, but is undoubtedly in the immediate area of CSUN loc. 153. The mollusks at CSUN loc. 153 must have undergone post-mortem transport from shallow-water sites into deep-water, submarine-fan paleoenvironments of the Tun- ca Canyon Formation. Several of the species found at CSUN loc. 153 or in the immediate vicinity (e.g., Anchu- ra phaba, Volutoderma magna, Crassatella elongata, Cymbophora triangulata, Glycymeris (Glycymerita) veat- chii, Pterotrigonia evansana, and Pinna sp.) were nor- mal-marine, shallow-depth dwellers (10 to 50 m) that also have been reported as transported remains in bathyal sub- marine-fan deposits of Campanian age in the Chatsworth Formation in the Simi Hills (Saul & Alderson, 1981). Etymology: The species is named for the state of Cali- fornia. Deussenia pacificana Squires & Saul, sp. nov. (Figures 10—12) Deussenia? n. sp. Saul, 1986:figs. 54—55. Diagnosis: A medium-shelled species of Deussenia with a subtabulate body whorl shoulder, bearing about 11 mod- erately strong axial ribs, and with prominent, closely spaced spiral ribs on body whorl. Description: Shell medium in size, 31 mm high (incom- plete); fusiform; spire of medium height, about two-fifths of total height of shell; spiral angle about 55°. Protoconch not preserved, and spire sculpture not preserved. Whorls about 4 1/2 (estimated). Upper part of body whorl con- stricted somewhat to broad, slightly concave ramp. Body whorl with about 11 (estimated) moderately strong and widely spaced axial ribs; strongest on shoulder and be- coming obsolete toward base of whorl. Body whorl with prominent, closely spaced spiral ribs persisting onto neck area. Aperture elongate-lenticular. Columella smooth. Dimensions of holotype: Height 31.5 mm, width 16 mm (incomplete at both extremities, especially the anterior end). Holotype: CASG 61598.01 [ex Stanford University 30031]. Type locality: CASG loc. 61598, latitude 120°55’40"N, longitude 35°43'45”"W. Distribution: Unnamed formation at Dip Creek, south shore of Lake Nacimiento, San Luis Obispo County, cen- tral California. Geologic age: Late Cretaceous (latest Maastrichtian) or possibly earliest Paleocene. Discussion: Only a single specimen has been found. It is small in size and could be a juvenile form. Taliaferro (1944) referred the Dip Creek strata to his “Dip Creek Formation,” but Durham (1968) referred to them as unnamed Upper Cretaceous and lower Tertiary rocks. At Dip Creek, the mollusks are shallow-marine forms that have undergone post-mortem transport and are within deep-water turbidites in beds of coarse-grained grit or conglomerate (Grove, 1986). More detailed geologic mapping is needed in the area before the Dip Creek sec- tion can be assigned to a formation. The outcrops along Dip Creek are usually covered by waters behind the Lake Nacimiento dam but are exposed during drought years (Squires & Saul, 1993). One can collect fossils along the nearby ridge top, but these specimens are harder to find and are more poorly preserved than those along the lake shore. The Dip Creek fauna contains some mollusks that re- semble genera or species usually considered to indicate a Cretaceous age, as well as some indicative of a Paleocene age. Taliaferro (1944) reported an unidentified ammonite from the fauna, and Saul (1983) reported a fragment of another ammonite, probably a Neophylloceras. Kirby & Saul (1995) reported a fragment of the bivalve Roudaria. These fossil remains suggest that at least the lower part of the section is of very Late Cretaceous age and that the upper half of the section, where the new species was col- lected, is no younger than earliest Paleocene. Based on specimens of Turritella peninsularis adelaidana (Metri- am, 1941) and 7. webbi (Saul, 1983), Saul (1983) as- signed the Dip Creek mollusks to a latest Maastrichtian and possibly an earliest Paleocene age. This age assign- ment was followed by Saul (1986) and Squires & Saul (1993). Etymology: The new species is named for the Pacific Ocean. Acknowlegments. Richard A. Flory (CSUC) donated two spec- imens of D. sierrana to the LACMIP collection. James W. Hag- gart (Geological Survey of Canada, Vancouver, British Colum- bia) facilitated our obtaining of specimens from the Pentz area. Lindsey T. Groves (LACMIP) provided access to the collections and loaned specimens. Jean DeMouthe (CAS) loaned specimens. Anton Oleinik (Department of Geology, Purdue University, In- diana) kindly shared his knowledge of buccinid gastropods. Mi- chael R. Cooper (Department of Geology, University of Durban- Westville, Durban, South Africa) graciously provided informa- tion on the type section of the Mzamba Formation, Pondoland, Transkei, South Africa. The manuscript benefited by reviews from David T. Dockery III (Mississippi Office of Geology) and an anonymous reviewer. Page 124 LITERATURE CITED AKERS, R. E. & T. J. AKERS. 1997. Texas Cretaceous Gastropods. 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Southern California Cretaceous formations and faunas with especial reference to the Simi Hills and Santa Monica Mountains. Pp. 15-29, pls. 1-3 in I. P. Col- burn & A. E. Fritsche (eds.), Pacific Section, Society of Eco- nomic Paleontologists and Mineralogists Fall Field Guide- book: Los Angeles, California. RENNIE, J. V. L. 1930. New Lamellibranchia and Gastropoda from the Upper Cretaceous of Pondoland (with an appendix on some species from the Cretaceous of Zululand). Annals of the South African Museum 28(pt. 12122):159—260, pls. 16-31. ROSENBERG, G. 1992. The Encyclopedia of Seashells. Dorset Press: New York. 224 pp. RusseLL, J. S., S. L. Baum & R. WATKINS. 1986. Late Coniacian to early Campanian clastic shelf deposits and molluscan as- semblages of the northeastern Sacramento Valley, Califor- nia. Pp. 179-196 in P. L. Abbott (ed.), Cretaceous Stratig- raphy Western North America. Pacific Section, Society of Economic Paleontologists and Mineralogists, Book 46: Los Angeles, California. SaAuL, L. R. 1983. Turritella zonation across the Cretaceous-Ter- tiary boundary, California. University of California Publi- cations in Geological Sciences 125:1—164, pls. 1-7. SaAuL, L. R. 1986. Mollusks of latest Cretaceous and Paleocene age, Lake Nacimiento, California. Pp. 25-31, figs. 1-60 in K. Grove & S. Graham (eds.), Geology of Upper Cretaceous and Lower Tertiary Rocks Near Lake Nacimiento, Califor- nia. Pacific Section, Society of Economic Paleontologists and Mineralogists, Volume 49: Los Angeles, California. SauL, L. R. 1988. New Late Cretaceous and Early Tertiary Per- issityidae (Gastropoda) from the Pacific slope of North America. Natural History Museum of Los Angeles County, Contributions in Science 400:1—25, figs. 1-128. SAUL, L. R. & J. M. ALDERSON. 1981. Late Cretaceous Mollusca of the Simi Hills: an introduction. Pp. 29—42, pls. 1-3 in M. H. Link, R. L. Squires & I. P. Colburn (eds.), Simi Hills R. L. Squires & L. R. Saul, 2000 Cretaceous Turbidites, Southern California. Pacific Section, Society of Economic Paleontologists and Mineralogists, Vol- ume and Guidebook: Los Angeles, California. SoHL, N. FE 1964. Neogastropoda, Opisthobranchia and Basom- matopohora from the Ripley, Owl Creek, and Prairie Bluff Formations. U.S. Geological Survey Professional Paper 331- B:1—344, pls. 19-52. Squires, R. L. & L. R. SAUL. 1993. A new species of Otostoma (Gastropoda: Neritidae) from near the Cretaceous/Tertiary boundary at Dip Creep, Lake Nacimiento, California. The Veliger 36(3):259—264, figs. 1-4. Squires, R. L. & L. R. SAUL. 1997. Late Cretaceous occurrences on the Pacific slope of North America of the melanopsid gastropod genus Boggsia Olsson, 1929. The Veliger 40(3): 193-202, figs. 1-17. STEPHENSON, L. W. 1941. The larger invertebrate fossils of the Navarro Group of Texas (exclusive of corals and crustaceans and exclusive of the fauna of the Escondido Formation). The University of Texas Publication 4101:1—641, pls. 1-95. TALIAFERRO, N. L. 1944. Cretaceous and Paleocene of Santa Lu- cia Range, California. Bulletin of the American Association of Petroleum Geologists 28(4):449—521. Ward, P. D., K. L. VERosuB & J. W. HAGGART. 1983. Marine magnetic anomaly 33—34 identified in the Upper Cretaceous of the Great Valley Sequence of California. Geology 11:90— OB. WENZ, W. 1943. Familia Galeodidae. Pp. 1211-1224, figs. 3452— 3479 in O. H. Schindewolf (ed.), Handbuch der Palazoolo- gie, Band 6, Prosobranchia, Teil 4. Gebriider Borntrager: Berlin. [reprinted 1960-1961] Woops, H. 1906. The Cretaceous fauna of Pondoland. Annals of the South African Museum 4(7):245—350, pls. 33-44. YERKES, R. F & R. H. CAMPBELL. 1979. Stratigraphic nomencla- ture of the central Santa Monica Mountains, Los Angeles County, California. U.S. Geological Survey Bulletin 1457E: E1-E31. YERKES, R. EF & R. H. CAMPBELL. 1980. Geologic map of east- central Santa Monica Mountains, Los Angeles County, Cal- ifornia. U.S. Geological Survey Map I-1146. APPENDIX LOCALITIES CITED CASG 61598. Dip Creek, NE 1/4 of section 30, T. 25 S, R. 10 E, U.S. Geological Survey Lime Mountain quad- rangle (7.5 minute, 1948, photorevised 1979), San Luis Obispo County, central California. Unnamed forma- Page 125 tion. Age: Late Cretaceous (latest Maastrichtian) or possibly earliest Paleocene. Collector: N. L. Taliaferro. CSUC PN31. At elevation of 530 ft., in a small canyon east of Highway 70, 716 m (2350 ft.) S and 69 m (225 ft.) E of NW corner of section 31, T. 21 N, R. 4 E, U.S. Geological Survey Cherokee quadrangle (7.5 min- ute, 1970), Butte County, northern California. Chico Formation, Pentz Road member (informal) of Russell et al. (1986). Age: Early Campanian. Collector: R. Watkins. CSUC PN32. At elevation of 530 ft., in a small canyon east of Highway 70, 754 m (2475 ft.) N and 107 m (350 ft.) W of SE corner of section 36, T. 21 N, R. 3 E, U.S. Geological Survey Cherokee quadrangle (7.5 minute, 1970), Butte County, northern California. Chi- co Formation, Pentz Road member (informal) of Rus- sell et al. (1986). Age: Early Campanian. Collector: R. Watkins. CSUN 153. [= LACMIP 11975.] At elevation of 1450 ft., in bottom of south fork of Garapito Creek, 533 m (1750 ft.) S and 521 m (1710 ft.) E of the intersection of the San Bernardino base line and Los Angeles City boundary, U.S. Geological Survey Topanga quadrangle (7.5 minute, 1952, photorevised, 1967), Santa Monica Mountains, Los Angeles County, southern California. Unnamed strata. Age: Late middle Campanian. Collec- tor: John Alderson, 1974—1987. [See Fritsche’s (1973) map for a plot of this locality on a detailed topographic base. | LACMIP 10833. Fossiliferous layers cropping out in beds of small gullies in field along Pentz Road [formerly Durham-Pentz Road], approximately 290 m (950 ft.) S and 107 m (350 ft.) E of NW corner of section 25, T. 1 N, R. 3 E, U.S. Geological Survey Cherokee quad- rangle (7.5 minute, 1970), Butte County, northern Cal- ifornia. Chico Formation, Pentz Road member (infor- mal) of Russell et al. (1986). Age: Early Campanian. Collector: W. P. Popenoe, 1931. LACMIP 27002. South fork of Garapito Creek, U.S. Geo- logical Survey Topanga quadrangle (7.5 minute, 1952, photorevised, 1967), Santa Monica Mountains, Los Angeles County, southern California. Unnamed strata. Age: Late middle Campanian. Collector: Mike Ham- ilton, circa middle 1970s. THE VELIGER © CMS, Inc., 2000 The Veliger 43(2):126-153 (April 3, 2000) Small, High-Spired Pulmonates from Mounts Mahermana, Iapiry, and Vasiha, Southeastern Madagascar, with Description of a New Genus, and with Conservation Statuses of 15 Streptaxid Species KENNETH C. EMBERTON Mollusk Department, Florida Museum of Natural History, Gainesville, Florida 32611-2035, USA AND TIMOTHY A. PEARCE! Delaware Museum of Natural History, Box 3937, Wilmington, Delaware 19807-0937, USA Abstract. Quantitative, replicated altitudinal transects on the three mountains yielded 20 small, high-spired pulmonate species in seven genera in three families. Descriptions are given of the two orculids Fauxulus andohahelae sp. nov. and F. gaillardi Fischer-Piette, Blanc, Blanc & Salvat, 1994; of the three subulinids Curvella vohimena sp. nov., Opeas tsiveryi sp. nov., and Subulina mamillata (Craven, 1880); and of the 15 streptaxids Gulella benjamini sp. nov., G. minuscula sp. nov., G. reeae sp. nov., Parvedentulina gen. nov. acutapex sp. nov., P. apicostriata sp. nov., P. esetra sp. nov., P. latembryohelix sp. nov., P. mahialamboensis sp. nov., P. margostriata sp. nov., P. ovatostoma sp. nov., P. rogeri sp. nov., Streptostele (Makrokonche) bougieformis sp. nov., S. (M.) latapex sp. nov., S. (M.) magnapex sp. nov., and S. (M.) vohimenensis sp. nov. Description of Parvedentulina gen. nov. includes three new combinations: P. glessi (Fischer-Piette, Blanc, Blanc, & Salvat, 1994) comb. nov.; P. metula (Crosse, 1881) comb. nov.; and P. simeni (Fischer- Piette, Blanc, Blanc, & Salvat, 1994) comb. nov. Streptaxids are a major component (17% on average) of the snail fauna in southeastern Madagascar, although not to the extent that they are in mainland-African rainforests (e.g. average 25%, maximum approx. 50% in eastern Tanzania). Distributional data indicate that, of the 15 streptaxid species, five are Critically Endangered and 10 are Endangered. INTRODUCTION Province. Mount Mahermana (Vohimena Chain) is north- east of the village of Esetra, Ilapiry (Vohimena Chain) is west of Mahialambo, and Vasiha (Anosy Chain) is west of Malio. Latitude and longitude are given in degrees, minutes, and seconds. MBI 373 (= Tol-1). Summit of Mt. Mahermana, 340 m, 24°26'12"S, 47°13'13"E. MBI 374 (= Tol-2). Slope of Mt. Mahermana, 300 m, 24°26'17"S, 47°13'10"E. MBI 375 (= Tol-3). Slope of Mt. Mahermana, 200 m, This paper is the second in a series of four that identify and describe the species reported on as morphospecies by Emberton et al. (1996, 1999) and Emberton (1997). This paper treats the Mahermana-Iapiry-Vasiha small, high- spired pulmonates; and it evaluates each streptaxid spe- cies for conservation status. METHODS And MATERIALS Collecting methods have been detailed by Emberton et 24°26'15"S, 47°13'04’E. al. (1996). Sixteen stations were collected and numbered MBI 376 (= Tol-4). Valley on Mt. Mahermana, 100 in the “Tol” series (for Tolagnaro = Fort Dauphin, the m, 24°26'22"S, 47°12'41’E. nearest city). These stations have been mapped by Em- MBI 377 (= Tol-5). Summit of Mt. Hapiry, 540 m, berton et al. (1996, 1999) and by Emberton (1997). To 24°51'40"S, 47°00'20’E. shorten the taxonomic descriptions, stations are described MBI 378 (= Tol-6). Ridge on Mt. Iapiry, 500 m, briefly below. Station numbers are in the series of the 24°51'33"S, 47°00'27’E. Molluscan Biodiversity Institute (MBI). All stations were MBI 379 (= Tol-7). Ridge, valley, and slope on Mt. restricted to primary forest that had no more than limited selective cutting. Ecological data are given by Emberton (1997:table 1). All stations are in Madagascar: Tulear Ilapiry, 400 m, 24°51'27"S, 47°00'38"E. MBI 380 (= Tol-8). Slope of Mt. Ilapiry, 300 m, 24°51'36"S, 47°00'40"E. MBI 381 (= Tol-9). Slope of Mt. Hlapiry, 200 m, 'To whom reprint requests should be sent. 24°51'39"S, 47°00'46’E. K. C. Emberton & T. A. Pearce, 2000 Table 1 Shell and reproductive characters used in descriptions. SHELL Diameter (0.1 mm) Height (0.1 mm) Height/Diameter Aperture mid-width (perpendicular to rotational axis)/shell diameter (0.1) 5. Aperture mid-height (parallel to rotational axis)/aperture width (0.1) 6. Width between upper and lower insertions of the peri- stome/aperture mid-width 7. Whorl number (0.1) 8. Coiling tightness (whorl number/In height) 9. Apex angle (degrees) 0) 1 gs IS . Spire angle (degrees) . Barrelling: outward departure from straight line of whorls between n-0.5 and approximate second whorl (% shell di- ameter) 12. Parietal and palatal dentition (no, yes) 13. Columellar dentition (no, yes) 14. First whorl diameter (mm) 15. Early coiling tightness (2/In diameter of first two whorls) 16. Apical sculpture 17. Transverse rib density (number in seventh or eighth 0.1 of body whorl) 18. Rib height (% shell diameter) 19. Diminishment of rib sculpture toward apex (%) 20. Shape of the suture (simple, shouldered, canaliculate) 21. Density of sutural notches (same as density of transverse ribs, *4 density of transverse ribs) 22. Umbilicus, regardless of columella (minute, imperforate or nearly so) 23. Columella reflection (slight, strong and flat, strong and rolled) 24. Sculpture other than transverse ribs REPRODUCTIVE SYSTEM 25. Penis length (mm) 26. Penis width (range in mm) 27. Penial sheath (incorporating loop of vas deferens) (yes, no) 28. Penial sheath height (mm) 29. Penial apical caecum (yes, no) 30. Apical caecum height (mm) 31. Penial general sculpture 32. Penial retractor muscle attachment point 33. Epiphallus (yes, no) 34. Vas deferens width(s) 35. Atrium approximate size 36. Vagina length 37. Spermathecal duct width 38. Spermatheca-plus-spermathecal-duct length 39. Oviduct contents (egg, embryo, nothing) 40. Oviducal-embryo whorl count (0.1) MBI 382 (= Tol-10). Lower summit of Mt. Vasiha, 860 m, 24°55'18”"S, 46°44’ 19’E. MBI 383 (= Tol-11). Slope of Mt. Vasiha, 700 m, 24°55'23”"S, 46°44’27’E. MBI 384 (= Tol-12). Slope of Mt. Vasiha, 500 m, 24°55'19"S, 46°44'45’E. Page 127 MBI 385 (= Tol-13). Valley on Mt. Vasiha, 400 m, 24°55'25"S, 46°44'45"E. MBI 386 (= Tol-14). Slope of Mt. Vasiha, 300 m, 24°55'37"S, 46°44'49’E. MBI 387 (= Tol-15). Slope of Mt. Vasiha, 200 m, 24°56'13"S, 46°45'13’E. MBI 388 (= Tol-16). Slope of Mt. Vasiha, 100 m, 24°56'20"S, 46°46'07’E. MBI 389 (= Tol-3-4). Incidental collecting between Tol-3 and Tol-4. MBI 390 (= Tol-1—2). Incidental collecting between Tol-1 and Tol-2. MBI 391 (= Tol-sub-5). Incidental collecting below summit of Mt. [lapiry, Tol-5. MBI 392 (= Tol-7-9). Incidental collecting between Tol-7 and Tol-9. Species identifications and comparisons were made us- ing Fischer-Piette et al. (1994) and Emberton (1994). For each species, the holotype or a representative shell was photographed in standard views. Orculids were pho- tographed at 16x magnification in apertural, side, and basal views; subulinids in apertural view at 6.4 or 25x and in subapical view at 40; and streptaxids in apertural view (6.4X, 10X, 16x, 25x, or 40X), in apical view at 40x, and—for Streptostele only—in side view showing penultimate and body whorls at 40 and sometimes in basal view at 40X (Figures 2-25). Twenty-four shell characters (Table 1, Figure 1) were measured, or measured and calculated, or scored from the photographs or from the shells themselves. At least one adult anatomy was available for eight spe- cies: one orculid, one subulinid, and six streptaxids. From each of these species, one or two reproductive systems were removed and photographed as they were turned and progressively dissected to expose characters. Sixteen re- productive-anatomical characters (Table 1, Figures 26— 35) were taken from the drawings or from the dissections themselves. Character matrices were prepared and used to code character-state data into the DELTA system (Partridge et al., 1993; Dallwitz et al., 1993), which was then used to generate natural-language species shell descriptions. The remainder of the descriptions were completed manually. For each streptaxid species, conservation status was evaluated using the new categories and criteria of the In- ternational Union for the Conservation of Nature and Natural Resources (IUCN, 1996). Ranges were estimated from distribution data in Emberton (in press). Rainforest extent and decline were assessed using Green & Sussman (1990), Sussman et al. (1994), and the most recently available topographic maps. SYSTEMATICS Higher classification follows Ponder & Lindberg (1997), Nordsieck (1986), and Vaught (1989). Type materials are fl NA _" l IN \\ Figure | Some shell features measured, scored, and used in calculating descriptive characters. 1WD - first whorl diameter; 2WD - first two whorls diameter; AA - angle at which apertural plane is inclined from rotational axis; AH - aperture height (inside dimension measured to and perpendicular to a line between columellar and upper peristome inser- tions); ApI - distance between the columellar and upper peristome insertions; ApO - amount of aperture occupied by previous whorl; AW - aperture width (inside dimension measured parallel to a line between the columellar and upper peristome insertions); AxA - apex angle; Ba - basal denticle; Br - barreling (outward departure from a straight line of a tangent to the whorls between n-0.5 and about the second whorl); C - columellar denticle; Co2 - second body whorl constriction; Con - body whorl constriction; D - shell diameter; FRb - final ribs near body whorl aperture; H - shell height; LC - lower columellar denticle; LP] - lower palatal denticle; MPI - middle palatal denticle; MPr - middle parietal denticle; OPr - outer parietal denticle; PCS - post-constrictional body whorl swelling; PeA - angle from greatest width of aperture plus peristome to rotational axis; Pel - peristome baso-palatal indentation (expressed as percent of basal peristome width, 1.e., to the unlabelled line above it in the figure); PH - aperture plus peristome greatest height as measured perpendicular to greatest width line; PI - palatal denticle; Pr - parietal denticle; PW 1 - aperture plus peristome width (measured parallel to aperture width); PW2 - aperture plus peristome greatest width (measured on Boucardicus Fischer-Piette & Bedoucha, parallel to or within 40 degrees of parietal-callus line); RbD - transverse rib density (number in estimated tenth of whorl); RC - recessed columellar denticle; RHr - rib hairs; S2D - swelling after second body whorl constriction; SA - spire angle; Su - suture depth one half whorl from aperture; UC - upper columellar denticle; Um - umbilicus size before any change in body whorl growth direction; UmF - final umbilicus total size; UPI - upper palatal denticle; Wh - whorl number. From Emberton & Pearce (1999). The Veliger, Vol. 43, No. K. C. Emberton & T. A. Pearce, 2000 Page 129 placed in the United States National Museum, Washing- ton, D.C. (USNM); temporarily in the Molluscan Biodi- versity Institute (MBI), whose collections will revert in the near future to the Florida Museum of Natural History; and in the Australian Museum, Sydney (AMS); the Mu- séum national d’Histoire naturelle, Paris (MNHN); and the Academy of Natural Sciences of Philadelphia (ANSP). For paratype localities, use the MBI catalog number to refer to the station numbers (in parentheses) above. MBI catalog numbers consist of station number, period, species number, D (dry) or A (alcohol-preserved), and when appropriate H (holotype) or P (paratype) or R (representative). Class GASTROPODA Clade HETEROBRANCHIA Clade PULMONATA Order STYLOMMATOPHORA Suborder ORTHURETHRA Superfamily CHONDRINOIDEA Family ORCULIDAE Genus Fauxulus Schaufuss, 1869 Fauxulus gaillardi Fischer-Piette, Blanc, Blanc, & Salvat, 1994 (Figures 2, 26, 36) Fauxulus sp. 1, Emberton et al., 1996:210. Emberton, 1997: 1148. Representative: MBI 382.03DR, Tol-10 (ad). Other specimens: MBI 382.03D (2 juv; AMS C.203437 [1 ad]), MBI 382.03A (1 ad), MBI 388.12A (1 ad [dis- sected]). Description of representative shell: Size and Shape. Shell sinistral. Diameter 3.7 mm; height 5.6 mm. Height-diameter ratio 1.5. Whorls 7.4. Spire angle 55 degrees. Apex angle 55 degrees. Spire profile convexity (outward departure from a straight line tangent to whorls n-0.5 and about the second whorl) 6% of shell diameter. Whorl periphery round. Suture depth one half whorl from aperture is 3% of shell diameter. Umbilicus before change in body whorl growth direction 3% of shell diameter. Final umbilicus 36% of shell di- ameter. Coiling tightness (whorl number divided by nat- ural logarithm of shell diameter) 5.7. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 44% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome in- sertions) 1.19. Distance between columellar and upper peristome insertions is 84% of aperture width. Penulti- mate whorl projecting into body whorl occupying 42% of aperture height measure. Columella not truncate. Col- umellar plica absent. Columella strongly reflected and flat. Apertural plane inclined downward 5 degrees from rotational axis. Aperture shape somewhat triangular. Peri- stome reflected; no second, internal peristome. Ratio of aperture width including peristome to aperture width ex- cluding peristome 1.4. Change in growth direction of body whorl occurs 0.4 whorls behind aperture. Apertural dentition present, typical of Fauxulus (two parietal, two columellar, one basal, and three palatal denticles). Middle parietal denticle height 42% of aperture width. Outer pa- rietal denticle height 35% of aperture width. Upper col- umellar denticle height 15% of aperture width. Lower columellar denticle height 15% of aperture width. Basal denticle height 7% of aperture width. Lower palatal den- ticle height 27% of aperture width. Middle palatal den- ticle form simple; height 11% of aperture width. Upper palatal denticle height 11% of aperture width. Apex. First whorl diameter 0.7 mm. First two whorls diameter 1.1 mm. Embryonic whorls smooth. Post-Embryonic Shell Sculpture and Color. Post-em- bryonic shell with crowded transverse ribs, about 15 in last tenth of body whorl; transverse ribs relatively straight above whorl periphery (angling forward down about 20 degrees), then at periphery, ribs bend abruptly back and inward (except not inward on the body whorl) then curve forward into the umbilicus; suture is a very short distance below the angled periphery, forming a canalicule. Basic shell color light yellow-brown. Peristome (excluding per- iostracum) white. Description of lower reproductive system (MBI 388.12A: 1 adult): Penis 2.9 mm long, slender but api- cally bulbous. Penis without sheath or caecum. Penial re- tractor muscle attached just below midpoint of penis (1.3 mm above its juncture with the atrium). Epiphallus 4.1 mm long, proximally slender (same width as penis), swol- len distally (to three times penis width) along two-fifths of its length. Vas deferens very slender along its entire length. Atrium small. Spermathecal duct and oviduct in- dependently entering the atrium, thus vagina absent. Sper- mathecal duct wide (0.3 mm) and long (4.9 mm, includ- ing spermatheca), internally lamellar, adherent to oviduct. Distribution: Known from Mts. Mahermana and Vasiha, 340 to 860 m elevation (this paper), and from another site in the Anosy chain at 1060 m elevation (Fischer-Piette et al., 1994). Fauxulus andohahelae Emberton & Pearce, sp. nov. (Figure 3) Fauxulus sp. 2, Emberton et al., 1996:210. Emberton, 1997: 1148. Page 130 The Veliger, Vol. 43, No. Figures 2—6 Figure 2. Fauxulus gaillardi Fischer-Piette, Blanc, Blanc & Salvat, 1994, Tol-10. Figure 3. Fauxulus andohahelae Emberton & Pearce, sp. nov., holotype. Figure 4. Subulina mamillata (Craven, 1880), Tol-12. Figure 5. Curvella vohimena Emberton & Pearce, sp. nov., holotype. Figure 6. Opeas tsiveryi Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 Holotype: USNM 860792 (ex MBI 383.01DH, Tol-11, ad). Paratypes: None. Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: southeast slope of Mount Vasiha, 700 m elevation: latitude 24°55’23’S, longitude 46°44'27’E: primary rainforest. Description of holotype shell: Shell Size and Shape. Shell sinistral. Diameter 3.3 mm; height 4.4 mm. Height-diameter ratio 1.3. Whorls 5.8. Spire angle 55 degrees. Apex angle 55 degrees. Spire profile convexity (outward departure from a straight line tangent to whorls n-0.5 and about the second whorl) 1% of shell diameter. Whorl periphery flattened. Suture depth one half whorl from aperture is 1% of shell diameter. Umbilicus before change in body whorl growth direction 5% of shell diameter. Final umbilicus 36% of shell di- ameter. Coiling tightness (whorl number divided by nat- ural logarithm of shell diameter) 4.9. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 42% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome in- sertions) 1.09. Distance between columellar and upper peristome insertions is 73% of aperture width. Penulti- mate whorl projecting into body whorl occupying 38% of aperture height measure. Columella not truncate. Col- umellar plica absent. Columella strongly reflected and flat. Apertural plane inclined downward 10 degrees from rotational axis. Aperture shape somewhat triangular. Peri- stome reflected; no second, internal peristome. Ratio of aperture width including peristome to aperture width ex- cluding peristome 1.4. Change in growth direction of body whorl occurs 0.4 whorls behind aperture. Apertural dentition present, typical of Fauxulus (two parietal, two columellar, one basal, and three palatal denticles). Middle parietal denticle height 25% of aperture width. Outer pa- rietal denticle height 20% of aperture width. Upper col- umellar denticle height 9% of aperture width. Lower col- umellar denticle height 16% of aperture width. Basal den- ticle height 6% of aperture width. Lower palatal denticle height 25% of aperture width. Middle palatal denticle split into two closely spaced bumps; height 7% of aper- ture width. Upper palatal denticle height 5% of aperture width. Apex. Embryonic whorls 1.7; diameter 0.4 mm. First whorl diameter 0.3 mm. First two whorls diameter 0.5 mm. Embryonic whorls smooth. Post-Embryonic Shell Sculpture and Color. Post em- bryonic shell with crowded transverse ribs, about 15 in last tenth of body whorl; transverse ribs relatively straight above whorl periphery (angling forward down about 20 degrees), then at periphery, ribs bend abruptly back and Page 131 inward (except not inward on the body whorl) then curve forward into the umbilicus; suture is a very short distance below the angled periphery, forming a canalicule; trans- verse ribs crossed by a few poorly defined weak spiral lines. Basic shell color orange-tan. Peristome (excluding periostracum) red-orange and white. Shell comparisons: Unique within the genus for its low spire, recessed lower palatal denticle, and nearly fused middle and upper palatal denticles. Distribution: Known only from Mt. Vasiha at 700 m elevation. Etymology: For the adjacent Andohahela Reserve. The type locality is expected to be incorporated into an even- tual Andohahela National Park. Suborder SIGMURETHRA Infraorder ACHATINIDA Superfamily ACHATINOIDEA Family SUBULINIDAE Genus Subulina Beck, 1837 Subulina mamillata (Craven, 1880) (Figures 4, 27, 37) Subulinidae sp. 1, Emberton et al., 1996:210. Emberton, 1997:1143, 1147, 1150, 1151. Representative: MBI 386.01DR, Tol-14 (ad). Other specimens: MBI 377.11D (1 ad, 1 juv), MBI 378.13D (2 ad), MBI 379.15D (2 ad, 4 juv), MBI 379.15A (1 juv), MBI 380.12D (2 juv), MBI 381.09D (4 ad, 2 juv), MBI 382.12D (2 juv), MBI 383.09D (1 juv), MBI 384.11D (7 ad, 15 juv), MBI 384.11A (1 ad, 2 juv), MBI 385.06D (21 ad, 30 juv), MBI 385.06A (1 ad, 2 juv), MBI 386.01D (17 ad, 52 juv; AMS C.203438 [1 ad]; MNHN [1 ad]; ANSP 400829 [1 ad]), MBI 386.01A (4 ad [2 dissected], 3 juv), MBI 387.06D (1 ad, 1 juv), MBI 388.04D (1 ad). Description of representative shell: Shell Size and Shape. Shell dextral. Diameter 4.1 mm; height 20.2 mm. Height-diameter ratio 5.0. Whorls 11.0. Spire angle 10 degrees. Apex angle 15 degrees. Spire profile convexity (outward departure from a straight line tangent to whorls n-0.5 and about the second whorl) 8% of shell diameter. Whorl periphery slightly flattened. Su- ture depth one half whorl from aperture is 5% of shell diameter. Final umbilicus 0% of shell diameter. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 7.8. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- Page 132 tions) 65% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome in- sertions) 0.85. Distance between columellar and upper peristome insertions is 88% of aperture width. Penulti- mate whorl projecting into body whorl occupying 6% of aperture height measure. Columella truncate. Columellar plica absent. Columella not reflected. Apertural plane in- clined downward; 15 degrees from rotational axis. Ap- erture shape ovate. Peristome simple; no second, internal peristome. No change in growth direction of body whorl near aperture. Apertural dentition absent. Apex. First whorl diameter 0.7 mm. First two whorls diameter 1.1 mm. Embryonic whorls eroded. Post-Embryonic Shell Sculpture and Color. Post em- bryonic shell with weak irregular growth ridges. Basic shell color yellow-tan. Shell variation: Occasional shells are more tightly coiled, with a more slender appearance, than the illus- trated representative. Shell comparisons: Whorls more flattened, sutures less crenulate, and slightly more slender than the introduced Subulina octona (Chemnitz). Body whorl occupying a greater proportion of the shell height than S$. manampet- saensis Fischer-Piette & Testud, as illustrated by Fischer- Piette et al. (1994:33). Description of lower reproductive system (MBI 386.01A: 2 adults): Penis 2.7 mm long, seemingly in two sections: base 1.8 mm long, slender and thin walled, api- cally swollen about a tongue-shaped verge that is rolled into a conical shape 0.5 mm long and, apically, 0.3 mm wide; penis apex (epiphallus?) 0.9 mm long, 0.1 mm wide basally, curved, swollen at the tip, with complex unde- termined internal structure. Penis lacks caecum and seems to lack sheath. Wall of basal penis smooth, without pi- lasters or hooks of any size. Penial retractor muscle at- tached at the penial apex. Epiphallus seemingly absent. Vas deferens slender along its entire length. Atrium small. Spermathecal duct joining oviduct 1.5 mm above the atri- um, hence vagina 1.5 mm long. Spermatheca and its duct 1.5 mm long, adherent to oviduct. Egg in oviduct 2.0 1.4 mm. Distribution: Mts. Hapiry and Vasiha from 100 to 700 m elevation (this paper), and from Rhodesia, the Trans- vaal, and widespread in Madagascar (Fischer-Piette et al., 1994). Comments: This species, probably introduced, seems to be an indicator of ecological degradation (Emberton, 1997). The Veliger, Vol. 43, No. 2 Genus Curvella Chaper, 1885 Curvella vohimena Emberton & Pearce, sp. nov. (Figure 5) Subulinidae sp. 2, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860793 (ex MBI 376.03DH, Tol-4, ad). Paratypes: MBI 379.16DP (1 juv; AMS C.203439 [1 ad]), MBI 380.26AP (1 ad), MBI 381.10D (1 ad, 2 juv), MBI 391.05AP (1 ad). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: valley on northwest slope of Mt. Mahermana, elevation 100 m: 24°26'22"S, 47°12'41"E: primary rainforest. Description of holotype shell: Shell Size and Shape. Shell dextral. Diameter 1.8 mm; height 3.9 mm. Height-diameter ratio 2.1. Whorls 4.8. Spire angle 30 degrees. Apex angle 35 degrees. Spire profile convexity (outward departure from a straight line tangent to whorls n-0.5 and about the second whorl) 2% of shell diameter. Whorl periphery round. Suture depth one half whorl from aperture is 7% of shell diameter. Final umbilicus 0% of shell diameter. Coiling tightness (whorl number divided by natural logarithm of shell di- ameter) 8.2. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 59% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome in- sertions) 0.89. Distance between columellar and upper peristome insertions is 70% of aperture width. Penulti- mate whorl projecting into body whorl occupying 2% of aperture height measure. Columella not truncate. Colu- mellar plica absent. Columella slightly reflected. Aper- tural plane inclined downward 5 degrees from rotational axis. Aperture shape ovate. Peristome simple; no second, internal peristome. No change in growth direction of body whorl near aperture. Apertural dentition absent. Apex. Embryonic whorls 1.6; diameter 0.7 mm. First whorl diameter 0.6 mm. First two whorls diameter 0.8 mm. Embryonic whorls with very weak spiral ridges plus sub-sutural crenulations. Post-Embryonic Shell Sculpture and Color. Post-em- bryonic shell with irregular growth ridges. Basic shell color pale yellow-tan. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Much smaller with fewer whorls than Curvella (?) poutiersi Fischer-Piette, Blanc, Blanc & K. C. Emberton & T. A. Pearce, 2000 Page 133 Salvat, 1994; upper edge of aperture angled upward less steeply so aperture is less high for its width. Distribution: Known only from the Vohimena-Chain Mts. Mahermana and I[lapiry, 100 to 400 m elevation. Comments: Assigned to Curvella because the upper ap- ertural edge curves backward at the upper suture. Etymology: For the Vohimena Mountain Chain, north of Fort Dauphin. Genus Opeas Albers, 1850 Opeas tsiveryi Emberton & Pearce, sp. nov. (Figures 6, 38) Subulinidae sp. 3, Emberton et al., 1996:210. Emberton, 1997:1147, 1150. Holotype: USNM 860794 (ex MBI 387.02DH, Tol-15, ad). Paratypes: MBI 385.07DP (1 ad), MBI 386.09DP (1 ad, 1 juv), MBI 387.02DP (2 ad, 2 juv; AMS C.203440 [1 ad]; MNHN [1 ad]; ANSP 400830 [1 ad]), MBI 387.02AP (3 ad [1 dissected]). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: eastsoutheast slope of Mount Vasiha, 200 m: 24°56'13"S, 46°45'13"E: primary rainforest. Description of holotype shell: Shell Size and Shape. Shell dextral. Diameter 1.8 mm; height 4.3 mm. Height-diameter ratio 2.4. Whorls 5.7. Spire angle 25 degrees. Apex angle 35 degrees. Spire profile convexity (outward departure from a straight line tangent to whorls n-0.5 and about the second whorl) 5% of shell diameter. Whorl periphery slightly flattened. Su- ture depth one half whorl from aperture is 6% of shell diameter. Final umbilicus 10% of shell diameter. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 9.7. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 59% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome in- sertions) 0.92. Distance between columellar and upper peristome insertions is 77% of aperture width. Penulti- mate whorl projecting into body whorl occupying 8% of aperture height measure. Columella not truncate. Colu- mellar plica absent. Columella slightly reflected. Aper- tural plane inclined downward 5 degrees from rotational axis. Aperture shape ovate. Peristome simple; no second, internal peristome. No change in growth direction of body whorl near aperture. Apertural dentition absent. Apex. First whorl diameter 0.5 mm. First two whorls diameter 0.8 mm. Embryonic whorls smooth then with transverse ribs. Post-Embryonic Shell Sculpture and Color. Post em- bryonic shell with strong, irregular transverse growth ridges, curved backward moderately strongly at upper su- ture. Basic shell color pale yellow-white. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Similar in shape to Opeas soulaianus Fischer-Piette & Testud but the translation rate is slower, leading to squatter individual whorls and more deeply impressed sutures; the adult height (4.5 mm) and number of whorls (5.5) are considerably less (10 mm and 9 whorls, respectively, in Opeas soulaianus). Description of lower reproductive system (MBI 387.02AP: 1 adult): Penis morphology unknown. Sper- mathecal duct joining oviduct 0.3 mm above the atrium, hence vagina 0.3 mm long. Spermatheca and its duct 0.4 mm long. Distribution: Known only from Mt. Vasiha, from 100 to 400 m elevation. Comments: A live-collected paratype similar in size to the holotype contained eggs, suggesting that this species is not simply a juvenile of the larger Opeas soulaianus Fischer-Piette & Testud, 1973. Further study is needed to determine whether this species is within the range of var- iation of O. soulaianus. Etymology: For our guide Tsivery (his name means “never lost’’?) of Malio, who bravely continued to collect snails on Mt. Vasiha after suffering a scorpion sting. Superfamily STREPTAXOIDEA Family STREPTAXIDAE Genus Streptostele Dohrn, 1866 Subgenus Streptostele (Makrokonche) Emberton, 1994 Streptostele (Makrokonche) magnapex Emberton & Pearce, sp. nov. (Figure 7) Streptaxidae sp. 1, Emberton et al., 1996:210. Emberton, 1997:1147. Emberton et al., 1999:table 2. Holotype: USNM 860795 (ex MBI 373.05DH, Tol-1, juv). Paratypes: MBI 374.13DP (1 juv), MBI 375.11DP (1 juv), MBI 377.12DP (1 juv), 378.14DP (1 juv), MBI 379.17DP (1 juv; AMS C.203441 [1 juv]), MBI The Veliger, Vol. 43, No. 2 Page 134 ee a’ Figures 7—8 Figure 7 (four views). Streptostele (Makrokonche) magnapex Emberton & Pearce, sp. nov., holotype. Figure 8 (four views). Streptostele (Makrokonche) vohimenensis Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 379.17AP (1 juv), MBI 382.13DP (1 juv), MBI 385.08DP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: summit of Mt. Mahermana, 340 m: 24°26'12”S, 47°13'13”E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 5.8 mm; height 11.7 mm. Height-diameter ratio 2.0. Whorls 4.7. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 1.9. Apex angle 130 degrees. Spire angle 30 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 4.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 2.0 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 1.6. Apical sculpture smooth. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 67% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 67% of aperture width. Penultimate whorl projecting into body whorl; occupying 6% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella strongly re- flected and rolled. Shell Sculpture. Transverse rib density 8 in the seventh or eighth tenth of body whorl; rib height 0.01% of shell diameter. Strength of rib sculpture completely diminished toward apex. Suture simple. Sutural notch density same as that of transverse ribs. Sculpture besides transverse ribs consists of evenly spaced spiral grooves, numbering 15 between sutures. Shell comparisons: Unique for its size, shape, and spiral- groove sculpture. Calls to mind a juvenile Streptostele manumbensis Emberton, 1994, but is about twice as large. Bears a superficial resemblance to a juvenile of the acavid Clavator obtusatus (Gmelin, 1791). Distribution and conservation status: Mts. Mahermana, Ilapiry and Vasiha, known from 200 to 860 m elevation. Also reported (Emberton, in press) from Pic St. Louis, 500-530 m. Thus apparently restricted to fragmented for- est patches within the Vohimena Chain and the southern Anosy Chain, within < 1000 km? of forest that is contin- ually declining in extent and in quality of habitat. Under IUCN (1996) criteria, this is an Endangered species. Comments: All specimens are juveniles. The adult could be the largest streptaxid known from Madagascar. Etymology: For the large (L. magni) apical whorls (L. apex). Page 135 Streptostele (Makrokonche) vohimenensis Emberton & Pearce, sp. nov. (Figure 8) Streptaxidae sp. 2, Emberton et al., 1996:210. Emberton, 1997:1147, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860796 (ex MBI 375.02DH, Tol-3, juv). Paratypes: MBI 373.25AP (1 juv), MBI 374.14DP (2 juv; AMS C.203442 [1 juv]), MBI 374.14AP (1 juv), MBI 375.02DP (1 juv; MHNH [1 juv]), MBI 379.18DP (1 juv), MBI 379.18AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: west slope of Mt. Mahermana, 200 m: 24°26'15"S, 47°13'04"E pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 5.3 mm; height 12.0 mm. Height-diameter ratio 2.3. Whorls 7.8. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 3.1. Apex angle 135 degrees. Spire angle 25 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 6.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 0.8 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 0.9. Apical sculpture smooth. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 51% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 74% of aperture width. Penultimate whorl projecting into body whorl; occupying 7% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 10 in the sev- enth or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture com- pletely diminished toward apex. Suture canaliculate. Su- tural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell comparisons: More tightly coiled than Streptostele manumbensis Emberton, 1994. Weaker sculpture and pro- portionately larger aperture than Parvedentulina metula (Crosse). Distribution and conservation status: Mts. Mahermana and Iapiry, 200 to 400 m elevation. Also found on Pics St. Louis and St. Jacques, southern Vohimena Chain; and, apparently, also on Mt. Sangasanga (430 m; 21°22'20"S, Page 136 The Veliger, Vol. 43, No. 2 47°52'05"E), near Kianjavato (Emberton, in press). Thus the known range of this species is in lowland rainforest from southwest of Mananjary, south to near Fort Dau- phin, an area of fragmented and rapidly disappearing for- est much less than 5000 km?. Under IUCN (1996) crite- ria, Streptostele vohimenensis sp. nov. is Endangered. Etymology: For the Vohimena Mountain chain north of Ft. Dauphin. Streptostele (Makrokonche) bougieformis Emberton & Pearce, sp. nov. (Figures 9, 10) Streptaxidae sp. 3, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860797 (ex MBI 384.02DH, Tol-12, juv). Paratypes: MBI 382.14DP (2 juv), MBI 383.16AP (1 juv), MBI 384.02DP (2 juv; AMS C.203443 [1 juv]), MBI 384.02AP (1 juv), MBI 385.09DP (1 juv), MBI 387.07DP (2 juv), MBI 387.07AP (1 juv), MBI 388.05DP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: east slope of Mt. Vasiha, 500 m: 24°55'19"S, 46°44'45”E: primary rain- forest. Description of holotype shell: Shell Size and Shape. Diameter 6.1 mm; height 16.1 mm. Height-diameter ratio 2.6. Whorls 8.9. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 3.2. Apex angle 140 degrees. Spire angle 20 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 3.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 0.9 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 0.8. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 55% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.8. Distance between columellar and upper peri- stome insertions is 78% of aperture width. Penultimate whorl projecting into body whorl; occupying 4% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Shell Sculpture. Transverse rib density 9 in the seventh or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture canaliculate. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell comparisons: More tightly coiled, more slender, and with a much greater constriction of the spire than Streptostele manumbensis. Also, the baso-columellar edge of the aperture is angular, as opposed to round. One- and-a-half times as high for the same number of whorls as Parvedentulina simeni (Fischer-Piette, Blanc, Blanc, & Salvat comb. nov.), but similar in shape. Distribution and conservation status: Known only from Mt. Vasiha, from 100 to 860 m elevation. Not found any- where else (Emberton, in press). A Critically Endangered species, because known from only one locality, with an extent of occurrence < 100 km’, and with its habitat con- tinually declining in area and quality (IUCN, 1996). Etymology: For its similarity in form (L. formis) to a candle (French bougie). Streptostele (Makrokonche) latapex Emberton & Pearce, sp. nov. (Figures 11, 28, 32, 39, 40) Streptaxidae sp. 4, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860798 (ex MBI 382.04DH, Tol-10, juv). Paratypes: MBI 377.13DP (1 juv), MBI 381.11DP (9 juv), MBI 381.11AP (1 juv), MBI 382.04DP (14 juv; AMS C.203444 [1 juv]; MNHN [1 juv]; ANSP 400831 [1 juv]), MBI 382.04AP (1 juv), MBI 384.12DP (1 juv), 387.13AP (1 ad [dissected], 1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: lower summit of Mt. Vasiha, 860 m: 24°55'18”"S, 46°44'19"E.: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 3.6 mm; height 6.7 mm. Height-diameter ratio 1.9. Whorls 5.6. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 2.9. Apex angle 150 degrees. Spire angle 25 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 6.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 1.2 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 0.9. Apical sculpture faint, complete trans- verse striae after first 1.1 smooth whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 58% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- K. C. Emberton & T. A. Pearce, 2000 Page 137 Figures 9-11 Figures 9-10. Streptostele (Makrokonche) bougieformis Emberton & Pearce, sp. nov.: holotype (Figure 9, two views) and paratype (Figure 10, three views). Figure 11. Streptostele (Makrokonche) latapex Emberton & Pearce, sp. nov., holotype. All scale bars | mm. Page 138 tions) 0.9. Distance between columellar and upper peri- stome insertions is 67% of aperture width. Penultimate whorl projecting into body whorl; occupying 6% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Shell Sculpture. Transverse rib density 7 in the seventh or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture shouldered. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell comparisons: Unique for its broadly domed apex and huge initial whorl. Description of lower reproductive system (MBI 387.13AP: 1 adult): Penis 2.7 mm long, 0.4—0.5 mm wide. Penial sheath present, 0.4 mm high, incorporating loop of vas deferens. Penial apical caecum absent. Penial general sculpture without pilasters, with an even field of medium-sized hooks, and without giant apical and basal hooks. Penial retractor muscle attached at the penial apex. Epiphallus absent. Vas deferens slender along its entire length. Atrium medium in size. Spermathecal duct joining oviduct 1.0 mm above the atrium, hence vagina 1.0 mm long. Spermathecal duct slender, at least 5 mm long, par- tially adherent to oviduct. Embryonic shells in oviduct 2.2 whorls. Distribution and conservation status: Mts. [lapiry and Vasiha, known from 200 to 860 m elevation. Not found anywhere else (Emberton, in press). Under IUCN (1996) criteria, an Endangered species, restricted to fragmented subpopulations within < 1000 km? of declining forest. Etymology: For the wide (L. /Jat-) apex (L. apex). Streptostele (Makrokonche) latembryohelix Emberton & Pearce, sp. nov. (Figure 12) Streptaxidae sp. 5, Emberton et al., 1996:210. Emberton, 1997:1147. Emberton et al., 1999:table 2. Holotype: USNM 860799 (ex MBI 381.04DH, Tol-9, juv). Paratypes: MBI 379.19DP (1 juv), MBI 381.04DP (0; AMS C.203445 [1 juv]). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Mahialambo: east- southeast slope of Mt. Hapiry, 200 m: 24°51’39"S, 47°00'46"E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 5.0 mm; height 8.4 mm. Height-diameter ratio 1.7. Whorls 6.4. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 3.0. Apex angle 155 degrees. Spire angle 35 de- The Veliger, Vol. 43, No. 2 grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 6.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 1.0 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 1.0. Apical sculpture strong, incomplete (su- tural) transverse striae after first 1.3 smooth whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 48% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 77% of aperture width. Penultimate whorl projecting into body whorl; occupying 4% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella strongly re- flected and flat. Shell Sculpture. Transverse rib density 9 in the seventh or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture shouldered. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell comparisons: Unique for its beehive shape and big initial whorl. Distribution and conservation status: Known only from Mt. Ilapiry, from only 200 to 400 m elevation. No report of any other localities (Emberton, in press). A Critically Endangered species: a single locality, declining habitat, extent < 100 km* (IUCN, 1996). Etymology: For the broad (L. /ati) embryonic whorl (L. helix). Genus Parvedentulina gen. nov. Parvedentulina Emberton & Pearce, gen. nov. Edentulina (?) Pfeiffer, 1856, Fischer-Piette et al., 1994:58— 62, figs. 39-43. Type species: P. ovatostoma sp. nov. Other species: P. acutapex sp. nov.; P. apicostriata sp. nov.; P. esetra sp. nov.; P. glessi (Fischer-Piette, Blanc, Blanc, & Salvat, 1994) comb. nov.; P. latembryohelix sp. nov.; P. mahialamboensis sp. nov.; P. margostriata sp. nov.; P. metula (Crosse, 1881) comb. nov.; P. rogeri sp. nov.; and P. simeni (Fischer-Piette, Blanc, Blanc, & Sal- vat, 1994) comb. nov. Description of shell: Shell Size and Shape. Diameter 1.6—4.0 mm; height 3.6-11.0 mm. Height-diameter ratio 2.1—2.8. Whorls 5.9— 7.5. Coiling tightness (whorl number divided by natural K. C. Emberton & T. A. Pearce, 2000 Page 139 Figures 12-15 Figure 12 (three views). Parvedentulina latembryohelix Emberton & Pearce, sp. nov., holotype. Figure 13. (four views) Parvedentulina rogeri Emberton & Pearce, sp. nov., holotype (tall shell and right apex) and paratype Tol- 9 (short shell and left apex). Figure 14. Gulella benjamini Emberton & Pearce, sp. nov., holotype. Figure 15. Gulella reeae Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. Page 140 The Veliger, Vol. 43, No. 2 logarithm of shell height) 3.5—4.8. Apex angle 100-150 degrees. Spire angle 15—25 degrees. Barreling (outward departure from a straight line of the whorls between n- 0.5 and about the second whorl) 7—10% of shell diameter. Umbilicus minute. Apex. First whorl diameter approximately 0.5—0.7 mm. Apical sculpture smooth; or faint, minute spiral striae; or faint to strong, complete or incomplete (sutural) trans- verse striae after the first 0.7—1.2 smooth whorl. Aperture. Aperture ovate in shape; width (inside di- mension, parallel to a line between the columellar and upper peristome insertions) approximately 0.4 shell di- ameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.8—1.0. Distance between columellar and upper peristome insertions is 0.7—0.9 aperture width. Penultimate whorl projecting into body whorl; occupying 4—10% of aperture height mea- sure. Parietal and palatal dentition absent; columellar den- tition absent. Outer lip unreflected. Columella reflection slight to strong and flat. Shell Sculpture. Transverse rib density 9—20 in the sev- enth or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture either not diminished or half diminished toward apex. Suture simple or shouldered. Sutural notch density either the same as or two-thirds the density of the transverse ribs. Besides transverse ribs, there may be faint spiral grooves below the whorl periphery. Description of lower reproductive system: Penis 0.5— 0.7 shell diameter. Penial sheath present, enclosing one- fifth to three-fourths of the length of the penis, and in- corporating a loop of the vas deferens. Penial apical cae- cum absent. Penial general sculpture without pilasters, with an even field of medium-sized hooks, and with one giant apical hook and one giant basal hook, each arising from a large papilla. Penial retractor muscle attached at the penial apex or just above it on the vas deferens. Epi- phallus absent. Vas deferens equally slender along its en- tre length or moderately wider alongside the penis. Atr- um small to medium in size. Spermathecal duct joining oviduct approx. 0.2 mm above the atrium, hence vagina approx. 0.2 mm long. Spermathecal duct slender and long; or broad, short, and undifferentiated from the sper- mathecal sac. Ovoviviparous; embryonic shell(s) in ovi- duct with 2.2 whorls. Comparisons: Shell. Differs from small Streptostele Dohrn, 1866 (sensu Pilsbry, 1919) in its conspicuously more domed and more bluntly apexed shell (compare Figures 16—24 [this paper] with Pilsbry’s [1919] plate XXI). Differs from Streptostele (Makrokonche) Emberton, 1994, in its much smaller adult size and slightly more domed shape. Differs from Edentulina Pfeiffer, 1856, in its much small- er size, greater delicacy, and unreflected outer lip. Differs from edentate Gulella in its taller, more capacious aper- ture, and less domed shell. Resembles no other known genera of Streptaxidae (Zilch, 1959-1960; Richardson, 1988). Reproductive Anatomy. Differs from all other known streptaxids in its gigantic basal and apical penial hooks. Comments: Streptostele (Makrokonche), because of its conchological similarities in most features except size, and genitalic similarity in all but giant penial hooks, may eventually need to be transferred to Parvedentulina gen. nov. or elevated to genus. Streptaxid relationships are poorly understood; any such revision should be based on synapomorphies rather than similarities. Etymology: For its partial resemblance to a small (L. parvi-) Edentulina Pfeiffer, 1856. Gender: feminine. Parvedentulina rogeri sp. nov. (Figures 13, 29, 33, 41, 42) Streptaxidae sp. 6, Emberton et al., 1996:210. Emberton, 1997:1143, 1146, 1149, 1151. Emberton et al., in press: table 2. Holotype: USNM 860800 (ex MBI 386.02DH, Tol-14, ad). Paratypes: MBI 378.22AP (1 juv), MBI 384.13DP (1 juv), MBI 385.10DP (2 ad, | juv), MBI 385.10AP (2 juv), MBI 386.02DP (9 ad, 44 juv; AMS C.203446 [1 ad]; MNHN [1 ad]; ANSP 400832 [1 ad]), MBI 386.02AP (3 ad [1 dissected], 12 juv), MBI 387.08DP (14 ad, 38 juv), MBI 387.08AP (4 ad, 16 juv), MBI 388.06DP (2 ad, 17 juv), MBI 388.06AP (1 ad, 8 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: southsoutheast slope of Mt. Vasiha, 300 m: 24°55'37"S, 46°44'49’E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 3.0 mm; height 6.4 mm. Height-diameter ratio 2.2. Whorls 7.0. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 3.8. Apex angle 150 degrees. Spire angle 25 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 10.0% of shell diameter. Umbilicus (irrespective of col- umella) minute. Apex. First whorl diameter 0.7 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 1.6. Apical sculpture strong, incomplete (su- tural) transverse striae after first 1.0 smooth whorl. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 49% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a K. C. Emberton & T. A. Pearce, 2000 Page 141 line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 74% of aperture width. Penultimate whorl projecting into body whorl; occupying 5% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella strongly re- flected and flat. Shell Sculpture. Transverse rib density 9 in the seventh or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture shouldered. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell variation: There is considerable variation in the size of the first whorl, as demonstrated in Figure 13. There is also some variation in post-embryonic coiling tightness, resulting in some shells appearing more slender (Figure 13). Shell comparisons: Similar in shape to Parvedentulina metula, but less than two-thirds as large for the same number of whorls. Description of lower reproductive system (MBI 386.02AP: 1 adult): Penis 2.1 mm long, 0.2-0.3 mm wide. Penial sheath present, 0.4 mm high, incorporating loop of vas deferens. Penial apical caecum absent. Penial general sculpture without pilasters, with an even field of medium-sized hooks, and with one giant apical hook and one giant basal hook, each arising from a large papilla. Penial retractor muscle attached at the penial apex. Epi- phallus absent. Vas deferens slender along its entire length. Atrium medium in size. Spermathecal duct joining oviduct 0.2 mm above the atrium, hence vagina 0.2 mm long. Spermathecal duct slender, at least 5 mm long, par- tially adherent to oviduct. Embryonic shells in oviduct 2.2 whorls. Distribution and conservation status: Mts. [lapiry and Vasiha, 100 to 500 m elevation. No other localities known (Emberton, in press). Thus apparently restricted to lower elevations of the southern Vohimena and Anosy Chains. An Endangered species: fragmented subpopulations with- in a continually diminishing extent of forest < 1000 km’. Etymology: For Roger Randalana, collector extraordi- naire, of Ambatolahy, near Ranomafana National Park. Parvedentulina acutapex Emberton & Pearce, sp. nov. (Figures 16, 30, 47) Streptaxidae sp. 9, Emberton et al., 1996:209, 210. Ember- ton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860801 (ex MBI 387.03DH, Tol-15, ad). Paratypes: MBI 385.11DP (4 ad, 8 juv; AMS C.203449 {1 ad]; MNHN [1 ad]; ANSP 400834 [1 ad]), MBI 385.11AP (3 ad [1 dissected], 2 juv), MBI 386.10DP (1 juv), MBI 387.03DP (6 ad, 3 juv), MBI 387.03AP (5 ad, 2 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: eastsoutheast slope of Mt. Vasiha, 200 m: 24°56'13"S, 46°45'13”E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 1.6 mm; height 3.9 mm. Height-diameter ratio 2.5. Whorls 6.0. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 4.4. Apex angle 100 degrees. Spire angle 20 de- grees. Barrelling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 8.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 0.6 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 6.5. Apical sculpture faint, complete trans- verse striae after first 1.1 smooth whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 51% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 1.0. Distance between columellar and upper peri- stome insertions is 75% of aperture width. Penultimate whorl projecting into body whorl; occupying 10% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 8 in the seventh or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture simple. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Unique in its acutely skewed initial whorl. Description of lower reproductive system (MBI 385.11 AP: 1 adult): Penis 0.8 mm long, 0.1 mm wide, swollen at apex. Penial sheath present, 0.6 mm high, cov- ering three-fourths of the length of the penis, and incor- porating a loop of the vas deferens. Penial apical caecum absent. Penial general sculpture unknown, but apparently without pilasters. Penial retractor muscle attached just above the penial apex on the vas deferens. Epiphallus seemingly absent. Vas deferens slender from the prostate to the sheath, swollen from the sheath to the penis. Atri- um small in size. Spermathecal duct joining the oviduct Page 142 The Veliger, Vol. 43, No. 2 Figures 16-21 Figure 16. Parvedentulina acutapex Emberton & Pearce, sp. nov., holotype. Figures 17, 18. Parvedentulina mar- gostriata Emberton & Pearce, sp. nov.: holotype (Figure 17) and paratype, Tol-3 (Figure 18). Figure 19. Parv- edentulina esetra Emberton & Pearce, sp. nov., holotype. Figure 20. Parvedentulina apicostriata Emberton & Pearce, sp. nov., holotype. Figure 21. Parvedentulina ovatostoma Emberton & Pearce, sp. nov., paratype, Tol-11. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 Page 143 0.2 mm above the atrium, hence the vagina is 0.2 mm in length. Spermatheca and duct conspicuously broad (0.1 mm) and short (0.5 mm), curved, with internal, longitu- dinal lamellar sculpture, the spermatheca not clearly de- marcated from its duct. Comments: The aberrant shell apex, spermatheca, penial sheath, and penial-retractor-muscle attachment suggest this may be a different (and new) genus. Penial sculpture remains unknown. Distribution and conservation status: Mt. Vasiha, 200 to 400 m elevation. Also reported from Pic St. Jacques (24°58'00"S, 46°57'50"E) at 170 m, but from nowhere else (Emberton, in press). Thus a lowland species, known from only two widely separated localities in the southern Anosy and Vohimena Chains, within < 1000 km? of de- clining forest. Thus an Endangered species (IUCN, 1996). Etymology: For the sharply (L. acut-) pointed spire (L. apex). Parvedentulina margostriata Emberton & Pearce, sp. nov. (Figures 17, 18) Streptaxidae sp. 10, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860802 (ex MBI 373.06DH, Tol-1, ad). Paratypes: MBI 373.06DP (1 ad, 3 juv), MBI 373.06AP (8 ad [1 dissected]), MBI 374.15DP (1 ad), MBI 374.AP (1 juv), MBI 375.12DP (1 ad; AMS C.203450 [1 ad]), MBI 375.12AP (1 juv), MBI 376.10DP (2 juv), MBI 376.10AP (1 ad). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: summit of Mt. Mahermana, 340 m: 24°26'12”S, 47°13'13"E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 1.6 mm; height 3.6 mm. Height-diameter ratio 2.3. Whorls 6.1. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 4.8. Apex angle 130 degrees. Spire angle 20 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 7.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl] diameter 0.5 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 11.0. Apical sculpture faint, complete trans- verse striae after first 1.2 smooth whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 54% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 1.0. Distance between columellar and upper peri- stome insertions is 69% of aperture width. Penultimate whorl projecting into body whorl; occupying 4% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 20 in the sev- enth or eighth tenth of body whorl; nb height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture simple. Sutural notch density same as that of transverse ribs. Sculpture besides transverse ribs consists of faint spiral grooves, numbering 6 below whorl periphery. Shell variation: Variation in coiling tightness makes some shells appear stouter than others: compare Figures 17 and 18. Shell comparisons: Similar to Parvedentulina rogeri sp. nov., in which six whorls give a shell height of approx. 4.9 mm, but with a height of only 3.2 mm for the same number of whorls. The first embryonic whorl is propor- tionately smaller. Thus this species is like a two-thirds miniature of P. rogeri. Description of lower reproductive system (MBI 373.06AP: 1 adult): Embryonic shell (whorl count not feasible) in oviduct. Anatomy otherwise unknown. Distribution and conservation status: Known only from Mt. Mahermana, 100 to 340 m elevation. No reports else- where, not even from adjacent mountains (Emberton, in press). Critically Endangered, by IUCN (1996) criteria: single location, extent < 100 km’, continuing decline in habitat. Etymology: For the fine striations (L. striat-) on the whorl periphery (L. margo). Parvedentulina esetra Emberton & Pearce, sp. nov. (Figure 19) Streptaxidae sp. 11, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860803 (ex MBI 374.01DH, Tol-2, ad). Paratypes: MBI 374.01AP (2 juv), MBI 375.13DP (1 juv), MBI 375.13AP (3 juv), MBI 376.11DP (1 juv), MBI 376.L1AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: WSW slope of Mt. Mahermana, 300 m: 24°26'17"S, 47°13'10"E: primary rainforest. Description of holotype shell: Page 144 Shell Size and Shape. Diameter 2.2 mm; height 4.6 mm. Height-diameter ratio 2.1. Whorls 5.9. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 3.9. Apex angle 130 degrees. Spire angle 25 de- grees. Barrelling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 8.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 0.7 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 3.1. Apical sculpture smooth. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 56% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 65% of aperture width. Penultimate whorl projecting into body whorl; occupying 6% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 12 in the sev- enth or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture simple. Sutural notch density two-thirds that of transverse ribs. No other sculp- ture besides transverse ribs. Shell comparisons: Very similar to Parvedentulina ro- geri sp. nov. in size and shape, but lacking that species’ distinctive apical sculpture, with a simple rather than shouldered suture, and with a less conspicuous but denser rib sculpture. Distribution and conservation status: As the above, an- other Critically Endangered species (IUCN [1996] crite- ria) known only from Mt. Mahermana, 100 to 300 m elevation, and nowhere else, even on adjacent peaks (Em- berton, in press). Etymology: For the village of Esetra. Parvedentulina apicostriata Emberton & Pearce, sp. nov. (Figure 20) Streptaxidae sp. 12, Emberton et al., 1996:210. Emberton, 1997:1146. Emberton et al., in 1999:table 2. Holotype: USNM 860804 (ex MBI 379.21DH, Tol-7, ad). Paratypes: MBI 379.21DP (1 ad, 2 juv; AMS C.203451 [1 juv]), MBI 379.21AP (1 juv), MBI 383.17AP (3 juv), MBI 385.17AP (1 juvy). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Mahialambo: ridge, The Veliger, Vol. 43, No. 2 valley, and slope on southsoutheast slope of Mt. [apiry, 400 m: 24°51°27"S, 47°00'38"E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 2.1 mm; height 4.2 mm. Height-diameter ratio 2.1. Whorls 6.2. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 4.3. Apex angle 135 degrees. Spire angle 25 de- grees. Barrelling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 8.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 0.5 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 4.6. Apical sculpture faint, minute spiral stri- ae. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 55% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 68% of aperture width. Penultimate whorl projecting into body whorl; occupying 6% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 11 in the sev- enth or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture simple. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell comparisons: Most similar to Parvedentulina ro- geri sp. nov. and Parvedentulina esetra sp. nov., but with much tighter coiling, a smaller first whorl, and apical spi- ral striae that are lacking in the others. Distribution and conservation status: Mts. llapiry and Vasiha, known from 400 to 700 m elevation. No other known localities (Emberton, in press). Another Endan- gered species, with fragmented subpopulations within < 1000 km? of declining forests in the southern Anosy and Vohimena Chains, thus meeting IUCN (1996) criteria. Etymology: For the apical. (L. apic-) spiral striae (L. Striat-). Parvedentulina ovatostoma Emberton & Pearce, sp. nov. (Figures 21, 22, 23, 48) Streptaxidae sp. 13, Emberton et al., 1996:209, 210. Ember- ton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860805 (ex MBI 386.04DH, Tol-14, ad). K. C. Emberton & T. A. Pearce, 2000 Page 145 Figures 22—25 Figures 22, 23. Parvedentulina ovatostoma Emberton & Pearce, sp. nov., paratype, Tol-9 (Figure 22), and holotype (Figure 23). Figure 24. Parvedentulina mahialamboensis Emberton & Pearce, sp. nov., holotype. Figure 25. Gulella minuscula Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. Paratypes: MBI 381.13DP (4 ad, 5 juv), MBI 381.13AP (2 ad, 4 juv), MBI 383.10DP (1 ad, 2 juv), MBI 383.10AP (3 ad, 1 juv), MBI 384.14DP (1 ad, 2 juv), MBI 385.12DP (1 ad, 2 juv; AMS C.203452 [1 ad]; MNHN [1 ad]; ANSP 400835 [1 ad]), MBI 385.12AP (2 ad [1 dissected], 6 juv), MBI 386.04DP (4 juv), MBI 388.08DP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: southsoutheast slope of Mt. Vasiha, 300 m: 24°55'37"S, 46°44'49"E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 1.7 mm; height 3.7 mm. Height-diameter ratio 2.2. Whorls 6.1. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 4.7. Apex angle 130 degrees. Spire angle 15 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 8.0% of shell diameter. Umbilicus (irrespective of colu- mella) minute. Apex. First whorl diameter 0.5 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 5.5. Apical sculpture strong, incomplete (su- tural) transverse striae after first 1.1 smooth whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 50% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 1.0. Distance between columellar and upper peri- stome insertions is 56% of aperture width. Penultimate whorl projecting into body whorl; occupying 4% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Page 146 Shell Sculpture. Transverse rib density 11 in the sev- enth or eighth tenth of body whorl; rib height 0.01% of shell diameter. Strength of rib sculpture diminishing by about half toward apex. Suture simple. Sutural notch den- sity same as that of transverse ribs. No other sculpture besides transverse ribs. Shell variation: There is great variation in the size of the first whorl and in coiling tightness, as demonstrated in Figures 21, 22, and 23. Shell comparisons: Very similar to Parvedentulina mar- gostriata sp. nov., but with looser embryonic coiling, slightly looser shell coiling, a conspicuously shorter cal- lus, rib sculpture only half as dense and diminishing to- ward the apex, and no spiral groove sculpture as in Par- vedentulina margostriata sp. nov. Description of lower reproductive system (MBI 385.12AP: 1 adult): Penis 0.9 mm long, 0.2-0.3 mm wide, but greatly constricted within sheath. Penial sheath present, 0.2 mm high, incorporating a loop of the vas deferens. Penial apical caecum absent. Penial general sculpture without pilasters, with a small but even field of medium-sized hooks, and with one giant apical hook and one giant basal hook. Penial retractor muscle attached at the penial apex. Epiphallus seemingly absent. Vas defer- ens slender at origin, insertion, and sheath, but swollen alongside the upper penis. Atrium size unknown. Sper- mathecal morphology unknown. Embryonic shell in ovi- duct 2.2 whorls. Distribution and conservation status: Mts. [lapiry and Vasiha, 100 to 700 m elevation. Other localities reported (Emberton, in press): Mt. Mahermana (300-340 m) and nearby Mts. Teloboko (530 m) and Esetra (summit); and, to the north, Miaranony. Thus the species seems to occur in fragmented subpopulations within < 5000 km? of de- clining forest in the southern Anosy Chain, throughout the Vohimena Chain, and north to Miaranony. Therefore, by IUCN (1996) criteria, this is an Endangered species. Etymology: For the ovate (L. ovat-) aperture (L. stoma, mouth). Parvedentulina mahialamboensis Emberton & Pearce, sp. nov. (Figure 24) Streptaxidae sp. 14, Emberton et al., 1996:210. Emberton, 1997:1147. Emberton et al., 1999:table 2. Holotype: USNM 860806 (ex MBI 379.02DH, Tol-7, ad). Paratypes: MBI 379.02DP (1 juv), MBI 379.02AP (1 juv), MBI 381.14DP (1 juv; AMS C.203453 [1 ad]), MBI 391.02DP (1 ad). Type locality: Madagascar: Tulear Province: northwest The Veliger, Vol. 43, No. 2 of Fort Dauphin: west of village of Mahialambo: ridge, valley, and slope on southsoutheast slope of Mt. Dapiry, 400 m: 24°51'27"S, 47°00'38’E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 3.1 mm; height 6.8 mm. Height-diameter ratio 2.2. Whorls 6.8. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 3.5. Apex angle 135 degrees. Spire angle 20 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 7.0% of shell diameter. Umbilicus (irrespective of colu- mella) imperforate or nearly so. Apex. First whorl diameter 0.8 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 1.6. Apical sculpture moderately strong, complete transverse ribs after first 0.7 whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 61% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.8. Distance between columellar and upper peri- stome insertions is 70% of aperture width. Penultimate whorl projecting into body whorl; occupying 5% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 16 in the sev- enth or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture simple. Sutural notch density two-thirds that of transverse ribs. No other sculp- ture besides transverse ribs. Shell comparisons: Similar to Parvedentulina glessi comb. nov. in size, shape, and sculpture, but with a con- spicuously more tightly coiled and more broadly domed apex, much weaker sculpture, and an essentially imper- forate umbilicus (perforate in P. glessi comb. nov.). Distribution and conservation status: Mt. Ilapiry, 200 to 400 m elevation. Emberton (in press) reports this spe- cies from Pic St. Jacques, the Midongy region, Manombo and environs, N of Manakara, and Kianjavato and envi- rons (total elevational range 50—660 m), but not from the northern Vohimena Chain or from anywhere in the Anosy Chain. Thus Parvedentulina mahialamboensis sp. nov. seems to occur in extremely fragmented subpopulations within windward rainforest from north of Fort Dauphin to Kianjavato. This forest is continually declining and is < 5000 km? in extent, so P. mahialamboensis meets IUCN (1996) criteria as an Endangered species. Etymology: For the village Mahialambo (east of Mount Ilapiry), whose name means “skinny pig.” K. C. Emberton & T. A. Pearce, 2000 Page 147 Genus Gulella Pfeiffer, 1856 Gulella reeae Emberton & Pearce, sp. nov. (Figures 15, 31, 34, 46) Streptaxidae sp. 7, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860807 (ex MBI 386.03DH, Tol-14, ad). Paratypes: MBI 381.12DP (1 ad), MBI 386.03DP (8 ad, 1 juv), MBI 386.03AP (2 ad), MBI 387.09DP (8 ad; AMS C.203447 [1 ad]; MNHN [1 ad]; ANSP 400833 [1 ad]), MBI 387.09AP (1 ad), MBI 388.07DP (2 ad, | juv), MBI 388.07AP (2 ad [1 dissected], 1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: southsoutheast slope of Mt. Vasiha, 300 m: 24°55'37"S, 46°44'49"E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 1.9 mm; height 4.3 mm. Height-diameter ratio 2.3. Whorls 6.5. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 4.5. Apex angle 135 degrees. Spire angle 20 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 11.0% of shell diameter. Umbilicus (irrespective of col- umella) minute. Apex. First whorl diameter 0.7 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 5.5. Apical sculpture smooth. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 53% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.7. Distance between columellar and upper peri- stome insertions is 90% of aperture width. Penultimate whorl not projecting into body whorl. Parietal and palatal dentition present; columellar dentition absent. Columella strongly reflected and flat. Shell Sculpture. Transverse rib density 7 in the seventh or eighth tenth of body whorl; rib height less than 0.005% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture shouldered. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Most similar to Gulella miaryi Fi- scher-Piette & Bedoucha, 1964, but with a much weaker parietal tooth, weaker sculpture, stouter shell, and—most importantly—shorter for more whorls. Description of lower reproductive system (MBI 388.07AP: 1 adult): Penis 1.1 mm long (not counting apical caecum), 0.2—0.3 mm wide at base and in upper third, bulging to 0.3 mm wide in lower two-thirds. Penial sheath absent, vas deferens free. Penial apical caecum present, 0.2 mm high. Penial general sculpture with two adjacent pilasters that run parallel in the upper half of the penis, then fuse to run as a single pilaster in the lower half; one of the upper pilasters bearing a large, tongue- shaped, pendant bulge. Penial wall otherwise without de- tectable sculpture. Penial retractor muscle attached at the penial apex and enveloping about half of the apical cae- cum. Epiphallus seemingly absent. Vas deferens broad at its departure from the prostate, then tapering and remain- ing slender until just before its entry into the penis, where it forms a bulblike swelling. Atrium medium in size. Spermathecal duct point of juncture with the oviduct un- certain. Spermathecal duct slender. Embryonic shell in oviduct 2.2 whorls. Distribution and conservation status: Mts. [lapiry and Vasiha, 100 to 300 m elevation. Also reported (Emberton, in press) far north at Miaranony (21°10'05’S, 47°33'20"E), near the eastern boundary of Ranomafana National Park, at 630 m, but nowhere in between. Even if the Miaranony identification is correct and this species ranges continuously from Mt. Vasiha to Miaranony, its range is still < 5000 km? of fragmented, declining forest, meeting criteria (IUCN, 1996) for Endangered status. Etymology: For Ruth Elizabeth Emberton, K.C.E.’s mother, born 13 July 1922. Gulella benjamini Emberton & Pearce, sp. nov. (Figures 14, 35, 43, 44, 46) Streptaxidae sp. 8, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860808 (ex MBI 381.05DH, Tol-9, ad). Paratypes: MBI 376.22AP (1 ad), MBI 377.14DP (2 ad), MBI 378.23AP (1 ad), MBI 379.20DP (1 ad; AMS C.203448 [1 ad]), MBI 379.20AP (3 ad [2 dissected]), MBI 380.13DP (1 ad), MBI 381.05DP (1 ad; MNHN [1 ad]). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Mahialambo: east- southeast slope of Mt. Hapiry, 200 m: 24°51'39"S, 47°00'46"E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 1.7 mm; height 3.2 mm. Height-diameter ratio 1.9. Whorls 6.0. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 5.2. Apex angle 125 degrees. Spire angle 35 de- grees. Barreling (outward departure from a straight line Page 148 The Veliger, Vol. 43, No. 2 Figures 26-35 Reproductive characters labeled on tracings from photographed (Figures 36—48) dissections. Figure 26. Fauxulus gaillardi. Figure 27. Subulina mamillata. Figure 28. Streptostele latapex Emberton & Pearce, sp. nov. Figure 29. Parvedentulina rogeri Emberton & Pearce, sp. nov. Figure 30. Parvedentulina acutapex Emberton & Pearce, sp. nov. Figure 31. Gulella reeae Emberton & Pearce, sp. nov. Figure 32. Streptostele latapex Emberton & Pearce, sp. nov. Figure 33. Parvedentulina rogeri Emberton & Pearce, sp. nov. Figure 34. Gulella reeae Emberton & Pearce, sp. nov. Figure 35. Gulella benjamini (bottom far right). Dissected penial tubes (Figures 32—35) show cut surfaces crosshatched. Abbreviations: A - albumen gland; B - bulge on penial pilaster; C - penial apical caecum; D - hermaphroditic duct; E - epiphallus; G - genital pore; H - penial sheath; k - hooks (penial sculpture); K - giant hook (penial sculpture); L1 - penial pilaster #1; L2 - penial pilaster #2; M - penial retractor muscle; P - penis; S - spermatheca or its duct; U - prostate-uterus (sometimes enclosing egg or embryos); V - vas deferens. K. C. Emberton & T. A. Pearce, 2000 Page 149 Figures 36—40 Pulmonate reproductive systems and penes. Figure 36 (two views). Fauxulus gaillardi Fischer-Piette, Blanc, Blanc & Salvat, 1994. Figure 37 (three views). Subulina mamillata (Craven, 1880). Figure 38 (one view). Opeas tsiveryi Emberton & Pearce, sp. nov. Figures 39—40 (six views total). Streptostele (Makrokonche) latapex Emberton & Pearce, sp. nov. All scale bars 1 mm. Page 150 The Veliger, Vol. 43, No. 2 Figures 41—45 Streptaxid reproductive systems and penes. Figures 41—42 (five views total). Parvedentulina rogeri Emberton & Pearce, sp. nov. Figures 43—45 (six views total). Gulella benjamini Emberton & Pearce, sp. nov. All scale bars | mm. of the whorls between n-0.5 and about the second whorl) 15.0% of shell diameter. Umbilicus (irrespective of col- umella) imperforate or nearly so. Apex. First whorl diameter 0.5 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 10.1. Apical sculpture faint, complete trans- verse striae after first 1.2 smooth whorls. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 42% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 0.9. Distance between columellar and upper peri- stome insertions is 86% of aperture width. Penultimate K. C. Emberton & T. A. Pearce, 2000 whorl not projecting into body whorl. Parietal and palatal dentition present; columellar dentition present. Columella strongly reflected and flat. Shell Sculpture. Transverse rib density 9 in the seventh or eighth tenth of body whorl; rib height 0.02% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture shouldered. Sutural notch density same as that of transverse ribs. No other sculpture besides trans- verse ribs. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Very similar to Gulella lubeti Fi- scher-Piette, Blanc, Blanc & Salvat, 1994, but with more tightly coiled whorls, two fewer whorls (six instead of eight), and with transverse ribbing sculpture beginning at 2.0 instead of at 2.7 whorls. Description of lower reproductive system (MBI 379.20AP: 2 adults): Penis 1.4 mm long (not counting apical caecum), 0.2—0.3 mm wide, tapered at apex. Penial sheath absent, vas deferens free. Penial apical caecum present, 0.3 mm high. Penial general sculpture with two adjacent, seemingly glandular pilasters that run parallel along the entire length of the penis; one of the penial- wall surfaces between them bears a sculpture of tiny chi- tinous hooks in regular array. The other penial-wall sur- face otherwise without detectable hook sculpture, but with faint latitudinal ridges. Penial retractor muscle at- tached at the penial apex and enveloping the entire apical caecum. Epiphallus seemingly absent. Vas deferens slen- der at origin and insertion and slightly swollen at mid- length. Atrium seemingly minute in size. Spermathecal duct joining the oviduct 0.6 mm above the atrium, hence the vagina is 0.6 mm in length. Spermathecal duct slen- der; length of duct plus spermatheca 1.6 mm. Distribution and conservation status: Known only from Mt. Mahermana and Mt. Ilapiry, known from 100 to 540 m elevation. No other localities reported, even from other mountains in the Vohimena Chain (Emberton, in press). An Endangered species, restricted to isolated subpopula- tions within the approx 500 km? extent of rapidly dimin- ishing Vohimena-Chain forests. Etymology: For Dr. Benjamin Andrianamahaja, National Director, Ranomafana National Park Project, in grateful appreciation for his assistance and moral support through- out this and other projects. Gulella minuscula Emberton & Pearce, sp. nov. (Figure 25) Streptaxidae sp. 15, Emberton et al., 1996:211. Emberton et al., 1999:table 2. Streptaxidae sp., Emberton, 1997:1140. Holotype: USNM 860809 (ex MBI 373.07DH, Tol-1, ad). Page ts Paratypes: MBI 376.23AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: summit of Mt. Mahermana, 340 m, 24°26'12”S, 47°13'13”E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 1.1 mm; height 2.4 mm. Height-diameter ratio 2.2. Whorls 5.0. Coiling tight- ness (whorl number divided by natural logarithm of shell height) 5.7. Apex angle 125 degrees. Spire angle 15 de- grees. Barreling (outward departure from a straight line of the whorls between n-0.5 and about the second whorl) 12.0% of shell diameter. Umbilicus (irrespective of col- umella) minute. Apex. First whorl diameter 0.5 mm. Early coiling tight- ness (2 divided by natural logarithm of diameter of first two whorls) 12.0 (much greater than). Apical sculpture smooth. Aperture. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) 50% of shell diameter; height-width ratio (inside dimensions, height measured to and perpendicular to a line between the columellar and upper peristome inser- tions) 1.0. Distance between columellar and upper peri- stome insertions is 76% of aperture width. Penultimate whorl projecting into body whorl; occupying 6% of ap- erture height measure. Parietal and palatal dentition ab- sent; columellar dentition absent. Columella slightly re- flected. Shell Sculpture. Transverse rib density 10 in the sev- enth or eighth tenth of body whorl; rib height 0.01% of shell diameter. Strength of rib sculpture not diminished toward apex. Suture simple. Sutural notch density same as that of transverse ribs. No other sculpture besides transverse ribs. Shell comparisons: Unique within the genus for its mi- nute size and narrowly protruding first whorl. Distribution and conservation status: Known only from the summit of Mt. Mahermana, 340 m elevation. Not re- ported anywhere else (Emberton, in press). As detailed for other such species above, Gulella minuscula sp. nov. meets IUCN (1996) criteria as a Critically Endangered species. Etymology: For the rather small (L.) size of the shell. STREPTAXID CONSERVATION STATUSES Analyses of individual species are given above in the spe- cies descriptions. To summarize, all 15 streptaxid species are proposed as either Endangered or Critically Endan- gered. The following five species should be listed as Crit- ically Endangered: Gulella minuscula sp. nov., Parved- entulina gen. nov. esetra sp. nov., P. margostriata sp. Page 152 The Veliger, Vol. 43, No. 2 Figures 46—48 Streptaxid reproductive systems and penes. Figure 46 (four views). Gulella reeae Emberton & Pearce, sp. nov. Figure 47 (three views). Parvedentulina acutapex Emberton & Pearce, sp. nov. Figure 48 (four views). Parved- entulina ovatostoma Emberton & Pearce, sp. nov. All scale bars 1 mm. nov., Streptostele (Makrokonche) bougieformis sp. nov., and S. (M.) latembryohelix sp. nov. The 10 species that should be listed as Endangered are: G. benjamini sp. nov., G. reeae sp. nov., P. acutapex sp. nov., P. apico- striata sp. nov., P. mahialamboensis sp. nov., P. ovatos- toma sp. nov., P. rogeri sp. nov., S. (M.) latapex sp. nov., S. (M.) magnapex sp. nov., and S. (M.) vohimenensis sp. nov. DISCUSSION These descriptions of 20 small, high-spired pulmonates provide systematic support for our previous distributional and ecological analyses of Mahermana-Ilapiry-Vasiha land snails (Emberton et al., 1996, 1999; Emberton, 1997). A previous paper (Emberton & Pearce, 1999) identified the 25 caenogastropods, described their 22 K. C. Emberton & T. A. Pearce, 2000 Page 153 small species, and analyzed their 17 species of Boucar- dicus Fischer-Piette & Bedoucha, 1965, for conservation statuses. In press are two additional papers describing the remaining Mahermana-Ilapiry-Vasiha pulmonates, and evaluating each charopid species for conservation status. Streptaxids are a major, previously underestimated component of the Madagascan land-snail fauna. Previous to this paper, Madagascar’s entire described streptaxid fauna numbered 30 species (Fischer-Piette et al., 1994; Emberton, 1994). Thus, collections from just three moun- tains, within a region representing less than 1% of Mad- agascar’s total area, have increased Madagascar’s de- scribed streptaxids by 50%. According to Emberton (in press:table 1), existing, identified collections from Mad- agascar include a total of 237 streptaxid morphospecies (far surpassing his earlier, pre-sorting estimate of 150 [Emberton & Rakotomalala, 1996]), and hundreds more species must survive in Madagascar that have never been collected. Of the 88 land-snail species found by us on Mts. Mah- ermana, Ilapiry, and Vasiha, 15 are streptaxids. Thus, on average, streptaxids constitute 17% of the fauna. This is a high proportion of carnivores, but falls short of the as- tonishing prevalence of streptaxids in eastern Tanzania, for example, where they averaged about 25%, and where at one rainforest site they composed about half the species and a third of the individuals (Emberton et al., 1997). Acknowledgments. We are grateful to the U.S. National Science Foundation and USAID for funding (grant DEB-9201060 to KCE); to staffs of the Ranomafana National Park Project, the Madagascar Département des Eaux et Foréts, and the Tolagnaro (Fort Dauphin) office of the World Wide Fund for Nature for logistical aid; to Roger Randalana and assistants from Esetra, Mahialambo, and Malio for collecting: to the late Felix Rako- tomalala for curatorial assistance; and to Lucia Emberton for help in mounting the photographs. LITERATURE CITED DaLLwitz, M. J., T. A. PAINE & E. J. ZURCHER. 1993. DELTA User’s Guide: A General System for Processing Taxonomic Descriptions. 4th ed. CSIRO Information Services: Mel- bourne, Australia. 136 pp. EMBERTON, K. C. 1994. Thirty new species of Madagascan land snails. Proceedings of the Academy of Natural Sciences of Philadelphia 145:147—189. EMBERTON, K. C. 1997. Diversities, distributions, and abundances of 80 species of minute-sized land snails in southeastern- most Madagascan rainforests, with a report that lowlands are richer than highlands in endemic and rare species. Biodi- versity and Conservation 6:1137—1154. EMBERTON, K. C. In press. A survey of Madagascar’s land mol- luscs: catalog of collections. Molluscan Biodiversity Insti- tute Occasional Publications 1:1—344. EMBERTON, K. C. & T. A. PEARCE. 1999. Land caenogastropods from Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with conservation statuses of 17 species of Boucardicus (Cyclophoridae). The Veliger 42(4):338—372. EMBERTON, K. C., T. A. PEARCE & R. RANDALANA. 1996. Quan- titatively sampling land-snail species richness in Madagas- can rainforests. Malacologia 38:203—212. EMBERTON, K. C., T. A. PEARCE & R. RANDALANA. 1999. Mol- luscan diversity in the unconserved Vohimena and the con- served Anosy mountain chains, southeast Madagascar. Bio- logical Conservation 89:183-188. FISCHER-PIETTE, E., C. P. BLANC, F BLANC & FE SALvart. 1994. Gastéropodes terrestres pulmonés. Faune de Madagascar 83: 1-552. GREEN, G. M. & R. W. SUSSMAN. 1990. Deforestation history of the eastern rain forests of Madagascar from satellite images. Science 248:212-215. Tucn. 1996. 1996 IUCN Red List of Threatened Animals. Inter- national Union for the Conservation of Nature and Natural Resources: Gland, Switzerland. 368 pp. Norpsieck, H. 1986. The system of the Stylommatophora (Gas- tropoda), with special regard to the systematic position of the Clausiliidae, Il. Importance of the shell and distribution. Archiv ftir Molluskenkunde 117:93-116. PARTRIDGE, T. R., M. J. DaLwitz & L. Watson. 1993. DELTA Primer: A General System for Processing Taxonomic De- scriptions. 4th ed. CSIRO Information Services: Melbourne, Australia. 15 pp. Pitssry, H. A. 1919. A review of the land mollusks of the Bel- gian Congo chiefly based on the collections of the American Museum Congo Expedition, 1909-1915. Bulletin of the American Museum of Natural History 40:1—370. PONDER, W. E & D. R. LINDBERG. 1997. Towards a phylogeny of gastropod molluscs: an analysis using morphological characters. Zoological Journal of the Linnean Society 119: 83-265. RICHARDSON, C. L. 1988. Streptaxacea: catalog of species, Part I, Streptaxidae. Tryonia 16:1—326. SUSSMAN, R. W., G. M. GREEN & L. K. SussMAN. 1994. Satellite imagery, human ecology, anthropology, and deforestation in Madagascar. Human Ecology 22:333-—354. VAUGHT, K. C. 1989. A Classification of the Living Mollusca. American Malacologists Inc.: Melbourne, Florida. 189 pp. ZitcH, A. 1959-1960. Gastropoda. Teil 2. Euthyneura. Band 6. Pp. 1-834 in O. H. Schindewolf (ed.), Handbuch der Palao- zoologie. Gebrtider Borntrager: Berlin. The Veliger 43(2):154—163 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 Three New Pacific Species of Halgerda (Opisthobranchia: Nudibranchia: Doridoidea) C. H. CARLSON aAnp P. J. HOFF' The Marine Laboratory, University of Guam, Mangilao, Guam 96923 Abstract. Three new species of Halgerda are described: H. batangas, a reticulate red-spotted species from the Philippines; H. okinawa a species from 53—76 m depth at Okinawa; and H. johnsonorum, a nocturnal H. willeyi-like species from the Marshall Islands with a relatively smooth dorsum. A specimen of H. willeyi Eliot, 1904, from Okinawa is described and illustrated for comparison with H. johnsonorum. INTRODUCTION Species of the genus Halgerda Bergh, 1880a (= Dictyo- doris Bergh, 1880b) are found throughout the tropical Indo-Pacific, as well as one in South Australia. Recent discussions of the genus can be found in Rudman (1978), Willan & Brodie (1989) and Carlson & Hoff (1993). Three new Halgerda species are presented in this paper, two from the Western Pacific and one much further east from the Marshall Islands. Because the reproductive systems are similar but dis- tinctly different among species, two figures are presented in order to show the distinctions. The first is a dorso- posterior view with the albumin-mucus gland pulled apart from the bursa copulatrix and prostate gland mass in or- der for the uterine duct and receptaculum to be seen. In most drawings the ampulla has been pulled down and away from the posterior of the mass and the receptaculum seminis is unburied. A second figure views the genital system from the right side. The radulae of the species described in this paper are typical of the genus in having very small teeth at the middle, gradually becoming larger toward the center of the half row and remaining large until near the end of the row. All are simply hamate with a flange on the inner edge of all except the outer laterals. Because the outer laterals may show distinction between species, they have been illustrated. SPECIES DESCRIPTIONS Halgerda okinawa Carlson & Hoff, sp. nov. (Figures 1—5) ?Halgerda graphica Basedow & Hedley, Eliot, 1913:12. Non H. graphica Basedow & Hedley, 1905. Halgerda sp. Gosliner, et al., 1996:160, fig. 563. Distribution: Halgerda okinawa has been recorded from ' Present address: P.O. Box 8019, Merizo, Guam 96916 Indonesia (Gosliner et al., 1996) and Okinawa (present study). Specimens: Seven specimens collected at Seragaki, Oki- nawa, Japan were furnished by Dr. Robert Bolland (all sizes are from the living animals). 110 mm, 67 m depth, 7 July 1996; 98 mm, 76 m, 22 July 1996, dissected; de- posited as paratype, Bishop Museum, Honolulu, BPBM 254044; 86 mm, 58 m, 14 January 1997; deposited as holotype, Bishop Museum, Honolulu, BPBM 253722; 66 mm, 73 m, 2 March 1997; deposited as paratype, Bishop Museum, Honolulu, BPBM 253723; 125+ mm and 114 mm, 53 m, 31 May 1997; 71 mm, 72 m, 10 April 1998. External morphology: The living animal (Figures 1, 2) is oblong-ovate with a mantle that lies along the substrate. The body is smooth, firm, and gelatinous. The dorsal sur- face has the typical ridges and depressions found in most Halgerda with the junctures sculptured with tubercles of varying height. The rhinophores arise out of a low smooth sheath, the branchia from a high smooth sheath. The rhinophore club is narrower than the base and has a slight posterior angulation. The branchia are divided into four main gills, the anterior having secondary branches and the posterior having two major branches with some secondary branching. The body is translucent white. The upper part of the tubercles is yellow which gradually diffuses into the white of the body. The yellow is internal and may or may not extend along the ridges between tubercles. Dark brown streaks of varying thickness line the tubercles. The streaks extend into the yellow of the tubercles though rarely is there any dark pigmentation on the tubercle tip. The streaks also are found along the ridges and reach the depressions between the ridges. Toward the mantle mar- gin the brown streaks become broken, often forming a series of spots. Brown spots may also appear irregularly on the body. The base of the rhinophores is translucent, the lamella yellow. The rhinophore sheath is unmarked except for a small tubercle of yellow postero-laterally on C. H. Carlson & P. J. Hoff, 2000 Page 155 Figures 1, 2 Halgerda okinawa Carlson & Hoff, sp. nov. Figure 1. Holotype (BPBM 253722). Length 86 mm. Seragaki, Oki- nawa, 58 m depth, 14 January 1997. Figure 2. Paratype (BPBM 253723). Length 66 mm. Seragaki, Okinawa, 73 m depth, 2 March 1997. Photos by Robert Bolland. most specimens. There is a posterior solid dark brown line the length of the rhinophores with some scattered dark brown spots on the sides of the stalk. The branchial sheath has a continuation of brown lines from the body as well as some touches of yellow. The branchia are translucent white with dark brown spots on the outer ra- chis and lines on the inner rachis. The protruding anus is white spotted with dark brown with a touch of yellow on the upper part. There is also spotting on the base of the branchia, sometimes being so concentrated that the base appears a solid dark brown. Morphological variations occur in terms of the number, size, and sculpturing of tubercles. The 66-mm paratype has a few high pointed tubercles, while in the 86-mm holotype they are low and rounded. Color variation oc- curs in terms of the number, length, and width of the brown streaks and number of lines on the inner surface of the branchia. The specimen illustrated from Indonesia by Gosliner et al. (1996:160) has yellow branchia as well as what appears to be a band of yellow on the mantle margin. One of the specimens from Okinawa had pale yellow branchia, two showed a touch of yellow on the posterior branchia, and one specimen had gold rather than yellow on tubercles and rhinophores. Specimens fixed in 10% formalin and preserved in 80% alcohol are translucent white with short dark brown streaks over the dorsum. Yellow was retained in only one specimen. The underside of the mantle is unmarked ex- cept for a few dark brown spots/lines at the juncture of mantle and body. The sides of the foot have some dark brown spots, in some cases forming incomplete rings. The genital pore was not marked. Internal morphology: The general internal arrangement is the same as for other species of Halgerda discussed by Rudman (1978) and Carlson & Hoff (1993). The visceral sac has a dusting of dark brown pigment posteriorly near the branchia; otherwise it is mostly transparent. The out- side of the oral tube was unpigmented; however, two dark brown spots were found inside. There was also some brown spotting inside the branchial pocket. With the vis- ceral sac open, the large bursa lies on and to the right of the midline. The esophagus and the aorta cross the top of the bursa. If the bursa is moved slightly away from the buccal bulb and to the right, the radular sac can be seen curling up and slightly to the left. The radula of the 98-mm dis- sected specimen was about 14 mm long and had a for- mula of 45 x 49.0.49 with the outer three laterals (Figure 3) reduced and simple. Within the reproductive system (Figures 4, 5), the am- pulla is long and folded. The uterine duct has a few folds and extends up under the vaginal duct where it enters the bursa copulatrix. The receptaculum seminis is peanut- shaped and is embedded in the prostate and albumin/mu- cus glands. The large bursa copulatrix is enfolded by, but not entirely covered by, the prostate gland. There is no thin layer of prostate over the bursa copulatrix. The pros- tate is brownish in color where it enfolds the bursa co- pulatrix and whitish as it folds around the base. The thin vas deferens exits from the heavy brownish prostate on top of the bursa copulatrix. It becomes larger and has one fold before reaching the large penial sheath. The vaginal duct joins the bursa copulatrix directly over the uterine duct. It widens only slightly at its distal end. Very large fleshy folds extend from the genital opening, through the body wall, creating the common genital vestibule. The Page 156 The Veliger, Vol. 43, No. 2 Figure 3 Halgerda okinawa Carlson & Hoff, sp. nov. Paratype (BPBM 254044). Length 98 mm. Radula-outer laterals. Scale = 100 jm. folds come together at the vaginal, mucus gland, and pe- nial openings. Discussion: Halgerda okinawa is the only species of Halgerda known to the authors that has a color other than the body color as an intrinsic part of the tubercles. Where- as most tubercle color in other species of Halgerda is made up of surface pigmentation which may extend a short distance into the tubercle, a section through the tu- bercles of H. okinawa reveals that the yellow pigment extends throughout the tubercle itself and diffuses into the translucent white of the body color. A similar pale yellow can be found on the ridges of many Halgerda willeyi Eliot, 1904, but the two species differ in terms of color pattern and dorsal morphology as well as in the genital system. Within the genital system, the almost bare bursa copulatrix, wrapped only in a narrow band of the prostate, separates H. okinawa from all other species of Halgerda that have been described. Eliot (1913:12) referred two animals from the ‘‘Oki- nawa Islands”? and Otaba to Halgerda graphica Basedow & Hedley, 1905. The specimens were described as having ridges with large tubercles at their junction, with both ridges and tubercles yellow. There were bold black lines and spots in the depressions. The rhinophore lamella was black, the lower part striped with black but showing a great deal of yellow. Outer lateral teeth were simple. The visceral sac was “‘blackish,”’ darker in one specimen than in another. Eliot’s description comes very close to H. oki- nawa, the differences being the black lamella on the rhi- nophores as well as the yellow “‘base” and the “‘black- ish” visceral sac. Etymology: The specific name okinawa refers to the lo- cale where the type specimens were collected. Figure 4 Halgerda okinawa Carlson & Hoff, sp. nov. Reproductive sys- tem: al, albumin gland; am, ampulla; be, bursa copulatrix; mu, mucous gland; pr, prostate; rs, receptaculum seminis; ud, uterine duct; vd, vas deferens: vgd, vaginal duct. Figure 5 Halgerda okinawa Carlson & Hoff, sp. nov. Reproductive sys- tem: am, ampulla; bc, bursa copulatrix; mo, mucous gland open- ing; mu, mucous gland; p, penial sheath; po, penial opening; pr, prostate; ud, uterine duct; vd, vas deferens; vgd, vaginal duct; vo, vaginal opening. C. H. Carlson & P. J. Hoff, 2000 Page 157 Figures 6, 7 Halgerda batangas Carlson & Hoff, sp. nov. Figure 6. Paratype (BPBM 254045). Length 35 mm. Anilao, Batangas, Luzon, Republic of the Philippines, 9 m depth, 23 April 1997. Figure 7. Length 40 mm. Dorsal anterior view to show reticulate markings. Anilao, Batangas, Luzon, Republic of the Philippines, 12 m depth, 22 April 1998. Halgerda batangas Carlson & Hoff, sp. nov. (Figures 6—11) Halgerda sp. Willan & Coleman, 1984:38, fig. 117. ?Halgerda sp. |. Tan et al., 1987:78, fig. 20. Halgerda sp. Colin & Arneson, 1995:180, fig. 833. Halgerda malesso Carlson & Hoff, Debelius, 1996:257. Non H. malesso Carlson & Hoff, 1993. Distribution: This species has been recorded from: Aus- tralia (Willan & Coleman, 1984); New Guinea (Colin & Arneson, 1995); the Philippines (present study; Debelius, 1996); and Indonesia (Debelius, 1996). Tan et al. (1987) mentioned an orange-lined Halgerda with orange-capped tubercles from Taiwan; the brief description and small color plate could possibly represent H. batangas. Material: Three specimens were collected from the Phil- ippines by Gary Williams, Marc Chamberlain, and Deb- bie Fugitt. Color transparencies of these specimens were made by the authors. One unmeasured specimen from Anilao (Batangas Province) collected in April, 1998, and computer scans of color transparencies of five specimens were made available by Mike Miller, San Diego, Califor- nia. (Sizes are from the living animals). 40 mm, Philip- pines, Cebu, Mactan Island, 7 m; 29 April 1997; depos- ited as holotype Bishop Museum, Honolulu, BPBM 253724; 35 mm, Philippines, Batangas, Anilao, 9 m; 23 April 1997; dissected; deposited as paratype, BPBM 254045; 40 mm, Philippines, Batangas, Anilao, 12 m; 22 April 1998. External morphology: The living animal (Figures 6, 7) is ovate with a smooth, firm gelatinous texture. A 40-mm specimen was 21 mm wide with a foot about 9 mm wide. Both ridges and tubercles are present, the major tubercles occurring at the juncture of the dorsal ridges. Other small- er tubercles may occur along the ridges or around the base of the major tubercles. Both rhinophores and branchia arise out of low, smooth sheaths. A small postero-lateral tubercle is present on the rhinophore sheaths of all but one of the specimens we were able to view. The rhinophore club is thinner than the base and has a slight posterior angulation. The apex is somewhat pointed. The branchia has four main gills. In the three specimens examined there was secondary branching in the anterior gills. One of the 40-mm speci- mens had two major branches in the posterior gills. The body is translucent white with a fine network of red-orange lines covering most of the dorsal surface. The tubercles are capped in red-orange surrounded by various intensities of white depending on the thickness of the dor- sal surface. The foot is lined with orange, and oral ten- tacles are tipped in orange. Rhinophores and branchia are translucent with sparse dark brown spots. The genital opening is not marked. Specimens fixed in 10% formalin and preserved in 80% alcohol have lost practically all of the red-orange color except for a couple of flecks on the tip of the largest tubercles. The brown on the rhinophores and branchia is retained as is the fine brown flecking on the visceral sac. Internal morphology: The visceral sac is dark brown with some clear areas. These clear areas match areas of dark brown on the inner body wall. We assume that pig- ment was transferred to the body wall as an artifact of preservation. The oral tube is unmarked. With the visceral sac opened, it can be seen that the bursa copulatrix is slightly to the left of the midline. The aorta, which is usually found passing over the top of the bursa, in H. batangas crosses the bursa along with the esophagus on Page 158 The Veliger, Vol. 43, No. 2 Figure 8 Halgerda batangas Carlson & Hoff, sp. nov. Paratype (BPBM 254045). Length 35 mm. Dorsal-dextral view: bce, bursa copu- latrix; bg, blood gland; bw, inner body wall; dg, digestive gland; es, esophagus; 1, intestine; mu, mucus gland; p, penis; s, stomach; v, vagina; vd, vas deferens; vgd, vaginal duct. the lower right side (Figure 8). The radular sac curls over and to the right. The radula of the 35-mm dissected specimen was a little under 5 mm long and had a formula of 59 < 52.0.52. The outer four teeth (Figure 9) were reduced in size with the outer three being flattened, the outermost rodlike, the penultimate bifid, and the antepenultimate irregularly bi- fid. The left side of the proximal end of the ribbon was folded over and lay across the right side. Within the reproductive system (Figures 10, 11) the ampulla is short and not convoluted. The uterine duct completes several loops before extending under the thin prostate gland covering the bursa copulatrix. A short tube from the uterine duct leads to the ovoid receptaculum seminis which is embedded in the gland mass between the bursa copulatrix, prostate, and albumin/mucus gland. The entire bursa copulatrix is covered by a thin layer of the prostate with a large prostatic mass folded under it. The vas deferens has one slight fold and terminates in a large penial sheath. The large vaginal duct exits from the thin layer of prostate over the bursa copulatrix. At the point at which it terminates, it is surrounded by a globular gland mass. The openings from the penis and mucus gland are adjacent to the vaginal opening within the gen- ital vestibule. Heavy folds line the common genital ves- tibule. These folds originate at the genital opening and Figure 9 Halgerda batangas Carlson & Hoff, sp. nov. Paratype (BPBM 254045). Length 35 mm. Radula-outer laterals. Scale = 10 pm. some extend into the vagina. Short brown streaks line some of the folds. Discussion: The fine network of lines on the dorsum and characteristics of the genital system distinguish Halgerda batangas from other species of Halgerda. Two other whitish species of Halgerda have been described that have a dorsal network of lines: Halgerda terramtuentis Bertsch & Johnson, 1982, and H. malesso Carlson & Hoff, 1993. Halgerda terramtuentis has relatively heavy gold lines, a submarginal gold line around the mantle, and white capped low tubercles. Halgerda malesso has fine orange lines that occasionally fuse to form patches of orange in the depressions, multiple fine submarginal orange lines around the mantle, and orange-capped, mod- erate to high tubercles. Red-capped tubercles as found in H. batangas are also found in H. carlsoni Rudman, 1978. Halgerda carlsoni is covered with fine red specks rather than a network of lines. The rounded glandular mass on the vaginal duct and the large penial sheath found in Halgerda batangas are similar to those shown by Willan & Brodie (1989:figs. 7, 8) for H. aurantiomaculata (Allan, 1932). They differ in that the glandular mass in H. aurantiomaculata is at the middle of the vaginal duct, whereas that in H. batangas is at the end of the duct. The penial sheath in H. aurantio- maculata is relatively larger than that of H. batangas. C. H. Carlson & P. J. Hoff, 2000 Figure 10 Halgerda batangas Carlson & Hoff, sp. nov. Reproductive sys- tem: al, albumin gland; am, ampulla; bc, bursa copulatrix; mu, mucous gland; pr, prostate; rs, receptaculum seminis; ud uterine duct; vgd, vaginal duct. Etymology: The specific name batangas is taken from the locale in the Philippines where this species is com- monly seen and where three of the specimens in the paper were collected. Halgerda johnsonorum, Carlson & Hoff, sp. nov. (Figures 12, 13, 15-17) Distribution: Halgerda johnsonorum has been recorded only from Kwajalein Atoll in the Marshall Islands. Material: Four specimens from the Ennubuy-Ennylabe- gan reef area of Kwajalein Atoll in the Marshall Islands were furnished by Scott and Jeanette Johnson. The spec- imens were found at night at a depth of 12 to 15 m; two in a small cave, and two in a surge channel ledge. (All sizes are from preserved material). 28 mm, 28 January 1989; deposited as holotype, Bishop Museum, Honolulu, BPBM 253725; 27 mm, 9 April 1993; opened dorsally; deposited as paratype, Bishop Museum, BPBM 253726; 20 mm, 15 October 1988; dissected; deposited as para- type, Bishop Museum, BPBM 254046; 25 mm, 2 July 1989; retained by authors; Halgerda willeyi Eliot, 1904. 98 mm (live measurement); Seragaki, Okinawa, 76 m; 22 July 1996; furnished by Dr. Robert Bolland; dissected for Page 159 imm Figure 11 Halgerda batangas Carlson & Hoff, sp. nov. Reproductive sys- tem: bc, bursa copulatrix; mo, mucous gland opening; mu, mu- cous gland; p, penial sheath; po, penial opening; pr, prostate; rs, receptaculum seminis; v, vagina (opened to show folds); vd, vas deferens; vgd, vaginal duct. comparative purposes. Deposited in Bishop Museum 254047. External morphology: Color transparencies of the living animals (Figures 12, 13) show that they are broadly ovate with a relatively low profile compared to other Halgerda. The mantle lies on the substrate. The surface has a texture like that of other Halgerda, but the overall body is some- what soft and flaccid, not the expected stiff and gelati- nous. There are low ridges on the dorsum with little, if any, sign of depressions between them. Their points of juncture are not marked by noticeable tubercles. The ridg- es that extend toward the mantle edge tend to become intermittent. Both rhinophores and branchia arise out of low smooth sheaths. The rhinophore club is narrower than the base and has a slight posterior angulation. The bran- chia is divided into four main gills with the posterior two being secondarily divided. The body varies from translucent white to translucent purplish gray. The ridges are yellow, becoming more in- tense at the points of juncture. The area between the ridg- es has dark brown and yellow lines. These lines form different shapes within the areas enclosed by the ridges but mid dorsally quite often run somewhat longitudinally. From the outer lateral ridges the lines radiate to the man- tle margin with the yellow lines becoming smaller, inter- Page 160 The Veliger, Vol. 43, No. 2 SAA GRE 3 os . : Figures 12-14 Halgerda johnsonorum Carlson & Hoff, sp. nov. Figure 12. Holotype (BPBM 253725). Length 28 mm (preserved). Kwajalein, Marshall Islands, 28 January 1989. Figure 13. Paratype (BPBM 253726). Length 27 mm (preserved). Kwajalein, Marshall Islands, 9 April 1993. (anterior mantle damaged) Photos by Scott Johnson. Figure 14. Halgerda willeyi, (BPBM 254047) Length 98 mm. Seragaki, Okinawa, 76 m. 22 July 1996. Photo by Robert Bolland. mittent, and rarely reaching the edge. The brown lines reach the margin, quite often branching before doing so. The branchia are translucent white with dark brown spots. The rhinophore sheath has a yellow margin; the branchial sheath is unmarked. The rhinophore base and lamella are translucent white with dark brown spots. A dark brown line marks the posterior of the base. In the preserved specimens the brown markings are retained; but the yellow is lost. A few lines on the un- derside of the mantle extend from the juncture of mantle and body wall to the edge of the mantle. The sides of the body have vertical brown lines and/or spots on the upper half and lines just above the foot. The top of the foot has numerous brown lines, some a continuation of the lines from the body. On some specimens brown lines appear on the bottom of the foot. A few brown lines lead into the genital pore. Internal morphology: The visceral sac is transparent with some sparse brown flecks. The general arrangement of the internal organs is typical of the genus. On the 27- mm specimen part of the whitish ampulla and the dark brown bursa copulatrix were covered by the prostate. The esophagus and aorta both crossed the top of the bursa. The oral tube is spotted in one specimen and unspotted in another. The radular sac curls up and to the right. The radula of the dissected specimen was about 5.5 mm long with a formula of 37 X 46.0.46. The outer six teeth (Fig- ure 15) are reduced in size with the outer three flattened and the penultimate bifid. Within the reproductive system (Figures 16, 17) the slightly coiled ampulla is short and most is embedded in the prostate gland. The uterine duct has one simple fold between its connection to the bursa copulatrix and the large albumin/mucus gland complex. The small ovoid re- C. H. Carlson & P. J. Hoff, 2000 Page 161 Figure 15 Halgerda johnsonorum Carlson & Hoff, sp. nov. Paratype (BPBM 254046). Length 20 mm (preserved). Radula-outer lat- erals. Scale = 100 pm. ceptaculum seminis is buried in the prostate gland. The bursa copulatrix is entirely covered by the large prostate gland with a large heavy part of the prostate extending around and to the base of the bursa copulatrix. The bursa copulatrix and albumin/mucus gland are relatively large. A thin vas deferens extends from the prostate gland on top of the bursa to the long, slightly widened penial sheath. The vaginal duct widens only slightly at the gen- ital vestibule. Folds at the genital opening extend through the body wall and line the common genital vestibule. Short brown lines appear externally and within the genital vestibule. Discussion: Halgerda johnsonorum differs from all other species of Halgerda the authors have examined in that it is somewhat soft and flaccid. In color, it is most similar to H. willeyi Eliot, 1904 (Figure 14) and could be mis- taken for that species if viewed only externally. Both have dark longitudinal lines mid-dorsally and dark lines radi- ating to the edge of the mantle margin. Both have dark lines under the mantle edges and along the top of the foot with the bottom of the foot being white. Both have yellow lining the ridges and deeper yellow at the juncture of the tubercles. In external coloration they differ slightly in that H. johnsonorum has brown spots and stripes along the sides of the body under the mantle, whereas H. willeyi has no markings in that area. Halgerda johnsonorum has brown spots over rhinophores and branchia, whereas H. willeyi has dark brown stripes on both. In external mor- phology they are distinct in that H. willeyi has very pro- nounced ridges and tubercles and, is stiff and gelatinous. The genital system for the H. willeyi, dissected for comparative purposes by the authors (Figures 18, 19), was as drawn by Eales (1938) except that the small gland Figure 16 Halgerda johnsonorum Carlson & Hoff, sp. nov. Reproductive system: al, albumin gland; am, ampulla; bc, bursa copulatrix; mu, mucous gland; pr, prostate; rs, receptaculum seminis; ud, uterine duct; vgd vaginal duct. Imm Figure 17 Halgerda johnsonorum Carlson & Hoff, sp. nov. Reproductive system: bc, bursa copulatrix; bw, body wall; go, genital opening; mu, mucous gland; p, penial sheath; pr, prostate; vd vas deferens; ved, vaginal duct. Page 162 Figures 18, 19 Halgerda willeyi, (BPBM 254047) Length 98 mm. Reproductive system: al-mu, albumin-mucus gland; am, ampulla; be, bursa co- pulatrix; go, genital opening (opened to show folds); po, penial opening; pr, prostate; rs, receptaculum seminis; ud, uterine duct; vd, vas deferens; vo, vaginal opening. The Veliger, Vol. 43, No. 2 oo : : \ Cay \ ENS \\ : Figure 20 Halgerda willeyi, (BPBM 254047) Length 98 mm. Radula-outer laterals. Scale = 100 pm. masses near the termination of the penial sheath were not as clearly defined as those shown by Eales. Halgerda wil- leyi differs significantly from H. johnsonorum in that the ampulla is large, long, convoluted, and not covered by the prostate gland. The uterine duct is sinuous and loops completely around the bursa copulatrix which is only par- tially covered by the prostate gland. The receptaculum seminis is comparatively larger than in H. johnsonorum. The penial sheath is short where it widens before entering the genital vestibule. Thin folds of tissue, rather than the larger heavy folds of H. johnsonorum, line the common genital vestibule. The radulae also differ in that H. willeyi has three sim- ple reduced outer laterals (Figure 20), whereas H. john- sonorum has six, the outer three flattened with the pen- ultimate being bifid. Etymology: The species is named for Scott and Jeanette Johnson who graciously furnished specimens and color transparencies. DISCUSSION The previously described Halgerda have had a smooth, stiff gelatinous texture. This is true of H. okinawa and H. batangas, but H. johnsonorum, while being smooth, is soft and flaccid with a thin body wall and dorsum. Rudman (1978) discussed a dark-colored visceral sac (fi- brous envelope) in four of the five species he studied. Of the three species described in this paper, H. batangas is dark while H. okinawa and H. johnsonorum are lightly pigmented. Willan & Brodie (1989) reported a lightly pigmented visceral sac for H. aurantiomaculata (Allan, 1932) as did Carlson & Hoff (1993) for H. guahan, H. malesso, and H. brunneomaculata. A major characteristic C. H. Carlson & P. J. Hoff, 2000 Page 163 of the Halgerda is a prostate-covered, or mostly covered, bursa copulatrix. Halgerda okinawa has a heavy band of the prostate that wraps over the bursa leaving both sides uncovered. This is the least coverage of the bursa copu- latrix of any of the described Halgerda. In addition to Halgerda batangas and H. aurantiomac- ulata, a glandular mass surrounding the vagina has also been reported for two other Halgerda: H. terramtuentis Bertsch & Johnson, 1982 (Kay & Young, 1969:fig. 28 [as H. sp. cf. graphica]) and H. malesso Carlson & Hoff (1993:fig. 15). These two species have large saclike va- ginas rather than the tubular forms found in H. auran- tiomaculata and H. batangas. Acknowledgments. This paper would not have been possible without the help of numerous people who furnished material. Specimens were furnished by Robert Bolland, Scott & Jeanette Johnson, Gary Williams, Marc Chamberlain, Michael Miller, and Debbie Fugitt. Color transparencies and/or computer scans were made available by Dr. Robert Bolland, Scott & Jeanette Johnson, and Michael Miller. We are also indebted to Linh Carlson for help with the Chinese translation. This paper is contribution No. 408 from the University of Guam Marine Laboratory. LITERATURE CITED BASEDow, H. & C. HEDLEY. 1905. South Australian nudibranchs, and an enumeration of the known Australian species. Trans- actions of the Royal Society of South Australia 20:34—60. BERGH, R. 1880a. Beitrage zue Kenntniss der japanischen Nu- dibranchien I. Verhandlungen der Zoologish-botanischen Gesellschaft in Wien 30:155—200. BERGH, R. 1880b. Malacologische Untersuchungen. in Reisen im Archipel der Philippinen von Dr. Carl Gottfried Semper. 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GosLIner, T. M., D. W. BEHRENS & G. C. WILLIAMS. 1996. Coral Reef Animals of the Indo-Pacific. Sea Challengers: Monte- rey. vit314 pp. Kay, E. A. & D. K. YounG. 1969. The Doridacea (Opisthobran- chia; Mollusca) of the Hawaiian Islands. Pacific Science 23(2):172—231. RUDMAN, W. B. 1978. The dorid opisthobranch genera Halgerda and Sclerodoris Eliot from the Indo-West Pacific. Zoological Journal of the Linnean Society 62(1):59-88. TAN, T.-H, J.-yY PAI & K.-c HsHA. 1987. An investigation on dis- tribution of nudibranch molluscs Along the coast Taiwan, R.O.C. Bulletin of Malacology, Republic of China 13:71— 90. WILLAN, R. C. & G. D. Bropie. 1989. The nudibranch Halgerda aurantiomaculata (Allan, 1932) (Doridoidea: Dorididae) in Fijian waters. The Veliger 32(1):69—80. WILLAN, R. C. & N. COLEMAN. 1984. Nudibranchs of Australasia. Australian Marine Photographic Index: Sydney. 56 pp. The Veliger 43(2):164—171 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 Taxonomic Revision of the Common Indo-West Pacific Nudibranch Phyllidia varicosa Lamarck, 1801 ALEXANDER FAHRNER Zoologische Staatssammlung Miinchen, Miinchhausenstr. 21, 81247 Miinchen, Germany (e-mail: kld1 130 @mail.Irz-muenchen.de) AND MICHAEL SCHRODL Zoologische Staatssammlung Miinchen, Miinchhausenstr. 21, 81247, Miinchen, Germany (e-mail: schroed] @ zi.biologie.uni-muenchen.de) Abstract. There has been a long-lasting debate on the taxonomic status of Phyllidia varicosa Lamarck, 1801, lacking a black foot stripe in preserved condition, and P. arabica Ehrenberg, 1831, retaining such a stripe after fixation. The present study demonstrates the significant influences of preservation on phyllidiid color patterns. Twenty-six P. varicosa specimens from Indonesia were fixed and stored under exactly the same conditions. While some specimens hardly faded at all, others completely lost their black pigmentation. Since living phyllidiids resembling P. varicosa all possess a foot stripe, its absence in a few preserved specimens described in the literature is easily explained as preservation artifacts. Still having faded remainders of a dark foot stripe, the recently rediscovered holotype of P. varicosa proves the synonymy of P. varicosa and P. arabica. In a detailed redescription, P. varicosa is shown to display considerable external and anatomical variability. Differences from Phyllidia alyta Yonow, 1996, from the Indian Ocean, of which type material has been reexamined, are critically discussed. INTRODUCTION The status of the very common and conspicuous Indo- West Pacific species Phyllidia varicosa Lamarck, 1801, has been subject to considerable dispute during recent years. The reason for all subsequent confusion was the description of a single specimen, on the basis of which Cuvier (1797) established the genus Phyllidia and which has been considered lost since 1866 (Willan et al., 1998). Lamarck (1801) introduced the specific name varicosa for the specimen described but not named by Cuvier. Both early authors in their descriptions neither mentioned a black longitudinal line on the foot sole nor did Cuvier (1804) show such a stripe in his drawings of the same specimen later (as Phyllidia trilineata Cuvier, 1804). On this basis, Yonow (1986, 1988, 1996) separated P. vari- cosa without a black foot stripe from the otherwise iden- tical Phyllidia arabica Ehrenberg, 1831, having a black stripe. In contrast, Brunckhorst (1993) considered P. arabica as a synonym of P. varicosa which was con- firmed by the recent rediscovery of the holotype (Willan et al., 1998). A number of very faint dark dashes on the foot sole was suspected to be the result of artificial fading. Doubting the possibility of extensive fading of the black pigment due to preservation, Yonow (1996) used the de- tailed appearance of the black foot stripe as a main char- acter to distinguish the new species Phyllidia alyta Yon- ow, 1996 from P. varicosa. The present study shows that some specimens of P. varicosa may fade so much due to preservatives that the black pigmentation on the dorsal and ventral surfaces, including the stripe on the foot sole, disappears complete- ly. Notes on the variability of the external morphology and a detailed anatomical description of P. varicosa are given. In addition, P. varicosa is compared with the sim- ilar species Phyllidia alyta. MATERIALS AnD METHODS A total of 27 specimens of P. varicosa ranging in length from 12 mm to 56 mm were examined. Twenty-six spec- imens were collected by A. Fahrner in September 1994 in Indonesian waters, at Lovina Beach/Bali (two speci- mens), Gili Trawangan/Lombok (22 specimens), and Gili Meno/Lombok (two specimens). Specimens were found, mostly by using SCUBA, on coral reefs in depths be- tween 2 and 21 m. They all were treated in exactly the same way and with the same chemicals: they were anes- thetized by slowly adding 20% MgCl, to the seawater, fixed in 10% buffered formalin and seawater for 24 hours, and preserved in 70% ethanol. One additional specimen from the Red Sea (Dahab, Egypt, October 1997, reef la- goon) was just fixed in 4% seawater buffered formalin. A. Fahrner & M. Schrédl, 2000 Twenty-five voucher-specimens were deposited in the Zoologische Staatssammlung Mtinchen (ZSM, Nos. 19983418—-32). The external morphology, especially the color patterns of the foot sole, and anatomical features of seven dissected specimens were investigated in detail un- der the binocular dissecting microscope. In addition, 14 specimens of Phyllidia alyta from the British Museum of Natural History (BMNH Nos. 1996107-8, 1996315—23), including the holotype and two paratypes, and a recently collected specimen from the Maldives (February 1999; ZSM No. 19991170) were studied. Finally, 102 preserved specimens of 17 other phyllidiid species, collected to- gether with the Indonesian P. varicosa specimens, were examined in order to compare the reaction of the color pigments to the preservatives in which the animals are stored. The central nervous system of one P. varicosa specimen was critical-point dried for scanning electron microscopy (SEM). SYSTEMATICS Phyllidiidae Rafinesque, 1814 Phyllidia Cuvier, 1797 Phyllidia varicosa Lamarck, 1801 Phyllidia varicosa Lamarck, 1801:66; Brunckhorst 1993:26— 29, figs. 2, 4-6, 23, 24, pl. 1 A-D. Phyllidia trilineata Cuvier, 1804:268, pl. A, figs. 1—6. Phyllidia arabica Ehrenberg, 1831:pages unnumbered. For an extensive synonymy of Phyllidia varicosa see Brunckhorst (1993), and slight modifications by Willan et al. (1998). External morphology (Figures 1A—F, 2A, B): The liv- ing and freshly preserved specimens from Indonesia in 1994 all fit precisely the description of P. varicosa given by Brunckhorst (1993), including the presence of a me- dian black stripe on the foot sole. Three blue-grey tuber- cle ridges between four longitudinal black lines and yel- low-capped tubercles characterize the dorsum of this spe- cies. In the examined specimens, the color patterns of the dorsum show a remarkable variability. The tubercle ridg- es may be continuous (Figure 1A) or broken (Figure 1C) and the four longitudinal black lines may be all connected (Figure 1C), all isolated (Figures 1A, 2A) or just two lines may touch (Figure 1E). Ventrally, the anterior edge of the foot is notched in 23 specimens and concave in three specimens. After four years of storage in ethanol, the Indonesian specimens differ considerably in the state of preservation of the color pigments (Table 1). Twelve specimens retain all the black and grey pigmentation both dorsally and ventrally (Figure 1C—F). They are hardly faded at all; only the yellow-orange of the tubercles and rhinophores is lost. Another six specimens still show a very distinct color pattern on the dorsal surface and also a distinct black stripe on the foot sole. However, the dark grey color Page 165 of the gills and the foot sole of these specimens gave way to a cream-white. Three specimens are more faded ven- trally and have lost parts of the foot stripe (Figure 2A, B). One specimen is very faded dorsally and ventrally, and the foot stripe is very indistinct. Finally, four speci- mens lack a line on the sole of the foot, and two of these specimens are so faded that all the black and grey col- oration dorsally and ventrally disappeared completely (Figure 1A, B). These two specimens already in life were lighter colored than the other collected P. varicosa spec- imens. Of the 22 preserved P. varicosa specimens retain- ing black markings on the foot sole, 12 are characterized by an unbroken, continuous line (Figure 1D, F), while 10 show different stages from a slightly broken to a distinct- ly broken or dotted stripe. The single P. varicosa specimen from the Red Sea dif- fers from the Indonesian specimens in having fewer black rays and marks on the mantle margin. The four longitu- dinal stripes on the dorsum are not connected, and the three tubercle ridges are continuous. It is remarkable that this specimen lost its black stripe on the foot sole com- pletely within less than 1 year of preservation, while the dorsal black pigmentation is hardly faded at all. Among 17 other phyllidiid species that have been in- vestigated within the framework of this study (see also Fahrner & Beck, in press), a total of eight specimens of Phyllidia elegans Bergh, 1869: Phyllidiella pustulosa (Cuvier, 1804); Phyllidiopsis striata Bergh, 1889; Phyl- lidiopsis annae Brunckhorst, 1993; and an undescribed species of Fryeria (Fahrner & Beck, in press), have lost all black pigmentation and faded entirely. Anatomy Digestive system (Figures 3A, B, 4): The foregut lacks any distinctive markings. A short, thin-walled oral tube is leading to the thick musculo-glandular pharyngeal bulb which has about the same length as width. Cream-colored bodies of the oral glands cover the postero-ventral, and in few specimens also the postero-lateral and postero-dor- sal parts of the pharyngeal bulb. Two retractor muscles arise posterodorsally on the pharyngeal bulb and insert the dorso-lateral body wall. These muscles are wide bands splitting into smaller bundles at their origin at the pharyngeal bulb. The pharynx shows considerable vari- ability in P. varicosa. After a short “‘intrabulbous”’ por- tion, a more or less swollen, tubular pharynx leaves the pharyngeal bulb posterodorsally, immediately forming a loop to the anterior-right in four specimens examined an- atomically (Figure 3A). In three specimens the pharynx also leaves the pharyngeal bulb posterodorsally but then extends posteriorly, turns and runs back anteriorly and turns a second time, thus forming an ‘‘S”’ in lateral view (Figure 3B). The esophagus narrows, passing through the central nerve ring (Figure 4), continues straight back- ward, and from anteriorly opens into the holohepatic, Page 166 The Veliger, Vol. 43, No. 2 A. Fahrner & M. Schrédl, 2000 Page 167 Table 1 Phyllidia varicosa: State of preservation of the color patterns of 26 specimens from Indonesia collected in 1994. Dorsal coloration (black and blue-grey) No. of speci- Entirely Entirely mens faded Faded Hardly faded faded 2 x x 1 x x 1 x 3 x 1 x x 6 x 12 x compact digestive gland, which occupies around two- thirds of the whole body cavity. No distinct stomach is detectable within the digestive gland; a caecum is absent. The intestine originates dorsally from the posterior third of the digestive gland and encircles the heart in a wide loop. The distal intestine runs posteriorly on the right side and ends medio-dorsally in the prominent anal papilla. The anal cone is situated in the center of a small cavity within a large tubercle, which dorsally has a round opening. Central nervous system (CNS; Figure 4): The CNS is positioned dorsally on the pharyngeal bulb. The cerebro- pleural ganglia are completely fused. The visceral loop is short, without distinct ganglia. The rhinophoral ganglia are separate, attached to the cerebral ganglion. The optic nerves are around two times as long as the diameter of the eyes. The pedal ganglia are situated next to the cer- ebropleural complex with the statocyst nestling in be- tween. The buccal ganglia are adjacent to the ventrolat- eral surface of the esophagus, posterior to the central nerve ring. A small gastroesophageal ganglion is inti- mately attached to each buccal ganglion. Reproductive system (Figure 5): The flat gonad overlies the anterior part of the digestive gland, being covered by the kidney dorsally and laterally. The triaulic anterior genitalia are situated in the space between the pharyngeal bulb and the digestive gland. The thin hermaphroditic duct passes into the spherical, brown colored ampulla which is faded to cream-white in some preserved speci- Ventral coloration (light and dark grey) Stripe on foot sole Faded Hardly faded Absent Faded Present x x x x x x x x x x x mens. The postampullar gonoduct is short, dividing into the vas deferens and oviduct, the latter entering into the female glands. The vas deferens enlarges into a convo- luted prostatic portion which extends to considerable length in larger specimens and distally narrows again into a muscular ejaculatory portion. The penis bears several rows of cuticular spines and enters into a common ves- tibulum with the vagina. The vagina is a narrow, rather long and straight duct which leads into the sperical, thin- walled bursa copulatrix. Next to the insertion, the vaginal duct arises from the bursa, bearing a stalked, muscular, elongate-ovate receptaculum seminis. The vaginal duct enters into the female gland mass close to the nidamental opening. Circulatory and excretory system: Like all phyllidiids, P. varicosa possesses secondary gills situated ventrolat- erally, in the groove between notum and foot. The flat, triangular shaped, grey gill leaflets are interrupted by the mouth anteriorly and the reproductive openings on the right side. Large and small gill leaflets alternate more or less regularly. The circulatory system corresponds to the description and drawings of Phyllidia flava Aradas, 1847 (as Phyllidia pulitzeri Pruvot-Fol, 1962) by Wagele (1984). The heart is placed mediodorsally on the digestive gland and kidney, in the posterior half of the body. Only two lateral vessels enter into the wide atrium. The mus- cular ventricle is situated anterior to the atrium. The large aorta runs to the blood gland which overlies the esopha- gus and parts of the reproductive organs. The blood gland Figure | Phyllidia varicosa, external variation of preserved specimens. A. Entirely faded specimen (29 mm; ZSM No. 19983418) with continuous tubercle ridges, dorsal view. B. Entirely faded specimen (29 mm; ZSM No. 19983418), ventral view. C. Hardly faded specimen (25 mm; ZSM No. 19983420) with broken tubercle ridges, dorsal view. D. Hardly faded specimen (25 mm; ZSM No. 19983420) with unbroken foot stripe, ventral view. E. Hardly faded juvenile specimen (13 mm; ZSM No. 19983421) with a single tubercle ridge, dorsal view. F Hardly faded juvenile specimen (13 mm; ZSM No. 19983421) with unbroken foot stripe. Scale bar = 1 cm. Page 168 The Veliger, Vol. 43, No. 2 Figure 2 Phyllidia varicosa, partly dissected specimen (53 mm; ZSM No. 19983422) showing a transitional stage of fading due to preservation. A. Dorsal view. B. Ventral view; the black foot stripe has disappeared except for some small remainders. Scale bar = 1 cm. : is a rounded to an elongate lobe which differs consider- ably in size. The syrinx is situated posteroventrally to the wide pericard, at the right side. The flat kidney covers the ovotestis dorsally and laterally. Mantle: The notum is thick and very tough. Basally, it is strengthened by a cross lamellar layer of strong, nee- dlelike spicules. From this layer, spicules arranged like bunches of flowers rise into each single tubercle; these structures are absent in areas between the tubercles. The spicules, reaching up to 1 mm in length and around 50 zm in diameter, are hollow, and all consist of calcium- carbonate. Silicate or chitinuous elements mentioned by Brunckhorst (1993) were not detected within the notum. Upper notum layers have a spongelike consistency and contain many large subepidermal glands. A. Fahrner & M. Schrédl, 2000 Page 169 DISCUSSION The 26 Indonesian specimens examined correspond to the description of P. varicosa by Brunckhorst (1993). These specimens, all fixed and preserved in the same way, show that even a complete loss of black pigmentation may be caused by preservation (see Table 1). Yonow (1996) stat- ed that she had “‘yet to see a P. arabica which is so faded that the black on the dorsal and ventral surfaces disap- pears completely’? and emphasized that preserved phyl- lidiid specimens retain their black coloration for very long periods. Most of the examined P. varicosa speci- mens after 4 years of preservation indeed still possess a very distinct black line on the foot sole (see Table 1, Figure 1D, F). However, four specimens lost this line, which had been present in life and in a freshly preserved state. Two of these specimens even faded completely and are entirely whitish now (see Figure 1A, B) due to pres- ervation artifacts. According to Yonow (1986, 1988, 1996), the only dif- ference between the two species P. varicosa and P. arab- ica is the absence vs. presence of a black stripe on the foot sole. Yonow had no example of a specimen without such a stripe, but based her theory of two separate species on Cuvier’s (1804) preserved holotype specimen of P. varicosa, considered lost for a long time. The rediscovery of this holotype in 1998 finally proved that P. varicosa has to be regarded as the valid name for the common Indo-West Pacific species with a longitudinal black foot stripe and that P. arabica is a junior synonym (Willan et al., 1998). Although collected more than 200 years ago, the holotype still possesses a line of very faded dark dashes on the foot sole. In the literature there are only three other descriptions of P. varicosa lacking a foot stripe (Gray, 1857; Quoy & Gaimard, 1832; Vayssiére, 1912; see Yonow, 1986), all referring to preserved spec- imens. In contrast, numerous authors mentioned speci- mens with a black foot stripe collected all over the Indo- Pacific region (see Brunckhorst, 1993; Yonow, 1986). The present study shows that it is inappropriate to use the black foot stripe to separate preserved phyllidiid spec- imens as it can fade partly or entirely, regardless of the state of fading of dorsal pigmentation (Table 1); in par- ticular, P. varicosa from the Red Sea lost the black foot Figure 3 Phyllidia varicosa, variability of the anterior digestive system. A. Large specimen (ZSM No. 19983422) with well-developed oral glands and a simple pharynx loop. B. Smaller specimen (ZSM No. 19983419) with less developed oral glands and a phar- ynx forming an “‘S” bend. Scale bars = 1 mm. Key: b, buccal ganglion; e, esophagus; g, gastoesophageal ganglion; og, oral glands; ot, oral tube; pb, pharyngeal bulb; ph, pharynx; rm, re- tractor muscles. Page 170 Figure 4 Phyllidia varicosa (ZSM No. 19983422), SEM micrograph of the CNS. Scale bar = 1 mm. Key: b, buccal ganglion; cpl, cere- bropleural ganglion; e, esophagus; p, pedal ganglion; ph, phar- ynx; r, rhinophoral ganglion; rn, rhinophoral nerve; y, eye. stripe in less than 1 year while retaining the black notum coloration. Phyllidia varicosa displays considerable individual variation in both external morphology and anatomy. This also refers to features used by Yonow (1996) to distin- guish the new species P. alyta Yonow, 1996, from P. varicosa. The black pigmentation on the foot sole of both P. varicosa and P. alyta does not always form a dotted line but is often an unbroken stripe (see Yonow, 1996: fig. 9 C, D; this study, Figure 1D, F). The four longitu- dinal black lines on the dorsum of P. varicosa do not always touch. There are also P. varicosa specimens with four individual lines or specimens with only the two inner lines being connected (this study). However, the recently collected specimen and the re-examined type and muse- um material from the Maldives clearly show the unique dorsal color pattern of P. alyta and highlight the differ- ences to P. varicosa: P. alyta completely lacks blue-grey coloration; its background color is white. The two inner of four black longitudinal stripes on the dorsum are al- ways connected by a short transverse line between the rhinophores (Yonow, 1996:fig. 9A; this study) which is absent in P. varicosa. Very few small black dots occur on the white mantle margin of P. alyta; black lines run- ning to the edge, which are typical for P. varicosa, are absent. The rounded tubercles are arranged in seven dis- tinctive rows (see Yonow, 1996:fig. 9A; this study), of which only the inner three form crests similar to those in P. varicosa. The genitalia of P. alyta generally agree with those of P. varicosa (see Figure 5). However, the bursa copulatrix of P. alyta possesses a distinct stalk, while the vagina and vaginal duct have a common insertion at the base of the bursa in P. varicosa (i.e., the bursa is not The Veliger, Vol. 43, No. 2 Figure 5 Phyllidia varicosa (ZSM No. 19983422), reproductive system. Scale bar = 1 mm. Key: am, ampulla; at, common atrium; bc, bursa copulatrix; fgm, female gland mass; hd, hermaphroditic duct; ov, oviduct; pr, prostatic vas deferens; rs, receptaculum seminis; v, vagina; va, vaginal duct; vd, muscular vas deferens. stalked) and, according to Brunckhorst (1993), in all other Phyllidia species. Moreover, seven dissected specimens of P. alyta, ranging from 21 mm to 27 mm preserved body length, are all sexually mature. In contrast, the fe- male gland mass is not yet developed in P. varicosa spec- imens smaller than 40 mm preserved body length. With a known living maximum length of 40 mm (Yonow, 1996), P. alyta is much smaller than P. varicosa, reaching up to 115 mm (Brunckhorst, 1993). Other anatomical features do not differ significantly. The pharyngeal bulb, according to Yonow (1996) sym- metrical in P. varicosa and asymmetrical in P. alyta, is variable in both species (Brunckhorst, 1993:fig. 4; this study) and highly influenced by the degree of develop- ment of oral glands. Bergh (1869) used foregut symmetry to contrast the “‘general symmetrical” foregut of the ge- nus Phyllidia from the long, highly folded, and therefore asymmetrically shaped foregut of Phyllidiella Bergh, 1869, but not to distinguish between different Phyllidia species. Within Phyllidia we agree with Marcus & Mar- cus (1970) and Brunckhorst (1993) in considering details of foregut symmetry an unreliable character. Showing re- markable intraspecific variability, the pharynx of P. var- icosa may be a rather narrow tube as claimed for P. var- icosa by Yonow (1996) or considerably swollen as in P. alyta. The pharynx either forms a large loop to the an- terior-right (Brunckhorst, 1993:figs., 4, 5; this study, Fig- ure 3A) or describes an ‘‘S”’ bend posteriorly (Brunck- horst, 1993:fig. 22; this study, Figure 3B) before passing the central nerve ring. The different conditions may re- A. Fahrner & M. Schrédl, 2000 flect different ontogenetic stages, since specimens with a pharynx forming an “S” were small and immature, while specimens with a pharynx-loop were all mature. At their insertion, the ribbon like pharyngeal bulb retractor mus- cles of P. varicosa may split into smaller bundles (this study, Figure 3), as described for P. alyta by Yonow (1996). The oral tube was not uniformly colored in the six P. alyta specimens anatomically examined by Yonow (1996); it was marked by black lines, a few black spots, or black pigmentation was completely absent as in P. var- icosa. In conclusion, P. varicosa and P. alyta cannot be dis- tinguished based on foregut anatomy or the appearance of the foot stripe, but with dorsal color pattern, arrange- ment of allosperm receptacles, and body size. Specimens from Mauritius assigned to P. alyta by Yonow (1996) differ from both the type material of P. alyta and from P. varicosa due to their dense notal tuberculation and dorsal color pattern; thus, their identity cannot be evalu- ated here. Acknowledgments. We are very grateful to Gerd Hagele, Moni Jocham, and Anette Meidert for their assistance in collecting Phyllidia varicosa and P. alyta. Joan Pickering and David Reid (BMNH) gave us museum material of P. alyta for re-examina- tion. Our gratitude goes to our Professors Gerhard Haszprunar (ZSM) and Horst Bohn (LMU) for kindly providing laboratory facilities in Munich. Special thanks are also directed to Teresa Saks (Munich) for language correction. Angel Valdés, Barry Roth, and an anonymous reviewer greatly contributed to improve the manuscript. LITERATURE CITED BerGu, L. S. R. 1869. Bidrag til en Monografi af Phillidierne. Naturhistorisk Tidsskrift (Kjobenhavn), Series B, 5:357— 542. BRUNCKHOorRST, D. J. 1993. The systematics and phylogeny of phyllidiid nudibranchs (Doridoidea). Records of the Austra- lian Museum, Supplement 16:1—107. Cuvier, G. L. C. F 1797. Sur un nouveau genre de mollusque. Bulletin des Sciences 1:105. Cuvier, G. L. C. F 1804. Mémoire sur la Phyllidie et sur le Page 171 Pleuro-branche, deux nouveaux genres de mollusques de Vordre des gastéropodes, et voisins des patelles les des os- cabrions, dont l’un est nu et dont l’autre porte une coquille cachée. 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Voyage de découvertes de |’ Astrolabe exécuté par ordre du Roi, pendant les années 1826-1829, sous le commandement de M. J. Dumont d’Urville. Tastu: Paris 2:1—320. VAYSSIERE, A. 1912. Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d’ Aden. Part 2. Annales du Faculté des Sciences de 1’ Université de Marseille, Supplement 20:5—157. WAGELE, H. 1984. Kiemen und Hamolymphkreislauf von Phyl- lidia pulitzeri (Gastropoda, Opisthobranchia, Doridacea). Zoomorphology 104:246—251. Wiitan, R. C., A. VaLpEs & D. J. BRUNCKHORST. 1998. No- menclatural implications from the rediscovery of the holo- type of Phyllidia varicosa Lamarck, 1801 (Nudibranchia: Phyllidiidae). Journal of Molluscan Studies 64:500—503. Yonow, N. 1986. Red Sea Phyllidiidae (Mollusca: Nudibranchia) with descriptions of new species. Journal of Natural History 20:1401—1428. Yonow, N. 1988. Red Sea Opisthobranchia 1: the family Phyl- lidiidae (Mollusca: Nudibranchia). Fauna of Saudi Arabia 9: 138-151. Yonow, N. 1996. Systematic revision of the family Phyllidiidae in the Indian Ocean Province: Part 1 (Opisthobranchia: Nu- dibranchia: Doridoidea). Journal of Conchyology, London 35:483-516. The Veliger 43(2):172-178 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 Redescription and Range Extension of Bathydoris aioca Marcus & Marcus, 1962 (Nudibranchia: Gnathodoridoidea) ANGEL VALDES anp HANS BERTSCH! Department of Invertebrate Zoology and Geology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA Abstract. Bathydoris aioca Marcus & Marcus, 1962, originally described from Baja California, is the only species of this genus known from the Pacific coast of North America. Examination of the original type material and newly collected specimens from Oregon, allows a redescription and range extension of this species. The digestive, reproductive, and central nervous systems are studied and re-illustrated, and scanning electron micrographs of the radula are presented for the first time. A comparison with other species of the genus leads to the conclusion that it constitutes a valid species. INTRODUCTION Bathydoris aioca is the only species of this genus known from the Pacific coast of North America. It was originally introduced by Marcus & Marcus (1962) on the basis of a single specimen collected from Baja California, Mexi- co, at 2753-2808 m depth. In the original description, details of the reproductive, digestive, and central nervous system morphology were overlooked. No additional spec- imens have been studied since the original description. Lance (1967) rediscovered the holotype of this species and deposited it in the collections of the Department of Invertebrate Zoology and Geology of the California Academy of Sciences (CASIZ). He cited this species un- der the incorrect spelling Bathydoris aoica. No further descriptions of the animal are given in that paper. The present paper redescribes B. aioca, based on a re- examination of the holotype and 12 additional specimens collected from Oregon, which constitute the second rec- ord of this species. SPECIES DESCRIPTION Bathydoris aioca Marcus & Marcus, 1962 (Figures 1—6) Type material: Holotype (by original designation): Northeast of Isla Guadalupe (29°40.2'—29°45.4'N, 117°06.6'—117°09.9'W), Baja California, Mexico, 2753— 2808 m depth, 15 February 1960, 71 mm _ preserved length, leg. R. Parker (CASIZ 018839). Lance (1967) mentioned the California Academy of Sciences type se- ries number for this species (306), which is no longer in use as the catalogue number. Additional material: Off Oregon coast (44°35.5'N, ' Mailing address: 192 Imperial Beach Blvd. # A, Imperial Beach, California 91932 125°35.3'W), USA, 2800 m depth, 29 April 1963, 1 spec- imen 47 mm preserved length, R/V Acona, cruise 6304 (CASIZ 115223). Off Oregon coast (44°44.5’N, 125°42'W), USA, 2850 m depth, 29 December 1963, 1 Figure 1 Bathydoris aioca, external morphology. A. Dorso-lateral view of a preserved specimen (CASIZ 115222), scale bar = 5 mm. B. Lateral view of a preserved specimen (CASIZ 115225), scale bar = 5 mm. A. Valdés & H. Bertsch, 2000 Page 173 Figure 2 Bathydoris aioca, external morphology. A. Dorsal view of a preserved specimen (CASIZ 113314), scale bar = 5 mm. B. Lateral view of a preserved specimen (CASIZ 113314), scale bar = 5 mm. C. View of the mouth area (CASIZ, 113314), scale bar = 2 mm. D. Branchial leaves arrangement (CASIZ 018939), scale bar = 5 mm. Abbreviations: as, anus; f, foot; gl, gill; 0, oral tentacle; r, renal opening; rh, rhinophore. specimen 23 mm preserved length, R/V Acona, cruise 6312 (CASIZ 115225). Off Oregon coast (44°44.8'N, 125°59.5'W), USA, 2800 m depth, 12 January 1965, 1 specimen 33 mm preserved length, R/V Acona, cruise 6501 (CASIZ 113315). Off Oregon coast (44°42.0'N, 125°37.3'W), USA, 2800 m depth, 20 February 1965, 1 specimen 23 mm preserved length, R/V Acona, cruise 6502 (CASIZ 113314). Off Oregon coast (44°39.2'N, 125°35.3'W), USA, 2810 m depth, 27 March 1966, 1 specimen 36 mm preserved length, R/V Yaquina, cruise 6503 (CASIZ 115224). Off Oregon coast (44°32.5'N, 125°24'W), USA, 2772 m depth, 9 April 1965, 1 speci- men 35 mm preserved length, R/V Yaquina, cruise 6504 (CASIZ 115226). Off Oregon coast (44°44.3'N, 125°41.4'W), USA, 2800 m depth, 24 October 1965, 1 specimen 15 mm preserved length, R/V Yaquina, cruise The Veliger, Vol. 43, No. 2 Figure 3 Bathydoris aioca, scanning electron micrographs. A. Rachidian and inner lateral teeth of the radula (CASIZ 018939), scale bar = 250 ym. B. Mid lateral teeth of the radula (CASIZ 018939), scale bar = 300 ym. C. Outer lateral teeth of the radula (CASIZ 018939), scale bar = 250 im. D. Tubercles and depressions of the mantle (CASIZ 018939), scale bar = 750 wm. A. Valdés & H. Bertsch, 2000 Figure 4 Bathydoris aioca. A. Jaw (CASIZ 113315), scale bar = 2 mm. B. Rhinophore of the holotype (CASIZ 018839), scale bar = 2 mm. C. Penis of the holotype (CASIZ 018839), scale bar = 5 mm. 6510 (CASIZ 115222). Locality and data unknown (prob- ably from the same area), 5 specimens 15-31 mm pre- served length (CASIZ 113316). External morphology: The color of the living animals is unknown. Preserved specimens are uniformly pale brown, lacking spots or traces of other colors. The body texture is soft, lacking spicules. In the dorsum, the teg- ument is very thin and the viscera are visible through the skin. The body is oval, elevated (Figures 1, 2A, B), with the rhinophores situated near the anterior edge of the no- tum. The gill is situated close to the posterior end of the notum. The dorsum is covered with small oval depres- sions (Figure 3D) which probably correspond to the place where the tubercles were attached in the living animal. Some minute, tentacular tubercles (Figures 1, 2A, B, 3D) remain in the laterals of the notum, anterior region of the body, and around the gill. The mantle margin is very narrow, only clearly visible in the anterior part of the body. The rhinophores are very elongate (Figure 4B). They have 45 large lamellae alternating with the same number of small ones in a 71 mm preserved length spec- imen. The gill consists of 18 bipinnate branchial leaves in a 71 mm preserved length specimen (Figure 2D). Each branchial leaf emerges from a depression in the mantle, Page 175 independently from the others. Very often the branchial leaves are arranged forming pairs, but they have been considered as two different leaves when they originate from different depressions in the surface of the mantle. The anus protrudes posteriorly (Figure 2D), closing the circle of branchial leaves. The renal pore also visibly pro- trudes (Figure 2D). The foot is strongly muscular and as long as the notum. It projects backward in a small tail. The oral tentacles are two long, flattened, lateral prolon- gations of the mouth area (Figure 2A—C). Anatomy: The buccal bulb consists of a large, oval mus- cular mass (Figure 5A). The jaws are two large, smooth plates (Figure 4A). They have thin growing marks crossed by radial lines. Except for that, there are no other marks or microsculpture. The radular formula is 64 X (81.1.81) in the holotype (71 mm long preserved speci- men) and 53 X (64.1.64) in a 33 mm preserved length specimen. The rachidian teeth are rectangular plates, lack- ing denticles (Figure 3A). The lateral teeth have a single elongate cusp, also lacking denticles (Figure 3B). The inner lateral teeth are very similar in shape to the mid laterals, but the cusp is shorter (Figure 3A). The outer lateral teeth are much smaller than the mid laterals, hav- ing also the same shape (Figure 3C). The two large, flat- tened salivary glands are connected to the laterals of the buccal bulb through two long ducts. The esophagus opens in the posterior end of the buccal bulb. It is short and wide, with the internal walls lined by strong ridges. These ridges are covered by a thin cuticule. Posteriorly, the esophagus opens into a muscular stomach, situated ven- trally in the viscera. Two ducts emerge at the end of the stomach (Figure 5C); one is the intestine, and the other is a short duct which opens into the digestive gland. The reproductive system is diaulic (Figure 5B). The ampulla is a long, very convoluted tube. It branches into a short oviduct and the prostate. The prostate is a long, highly convoluted undifferentiated duct. It expands dis- tally into the muscular ejaculatory portion. The vaginal duct is wide and long. It runs inside of the female gland most of its length. Near its distal end the vaginal duct connects with the bursa copulatrix. The penis is long and smooth, lacking hooks (Figure 4C). In the central nervous system (Figure 6) the cerebral and pleural ganglia are separated and also distinct from the pedal ganglia. The cerebral ganglia are very large and they appear to be divided in two regions. From each ce- rebral ganglion emerge five nerves. No visceral loop has been observed. At the end of the optic nerves no eyes have been observed though careful examination, and this species is probably blind. The rhinophoral ganglia are sit- uated in the middle of the cerebral ganglia. The pleural ganglia are small; from each one emerge seven nerves. The pedal ganglia are twice as large as the pleural. From each one emerge three nerves, the pedal and parapedal commissures. Also, from the right pleural ganglion Page 176 The Veliger, Vol. 43, No. 2 Figure 5 Bathydoris aioca, anatomy. A. Dorsal view of the internal organs (CASIZ 113316), scale bar = 2 mm. B. Repro- ductive system (CASIZ 113315), scale bar = 1 mm. C. Dissected digestive tract (CASIZ 113316), scale bar = 2 mm. Abbreviations: a, ampulla; b, blood gland; bc, bursa copulatrix; bm, buccal mass; d, digestive gland; e, esophagus; fg, female gland; h, heart; i, intestine; pe, penis; pr, prostate; rs, renal sac; s, salivary gland; st, stomach; v, vaginal duct. emerges the genital nerve. The buccal ganglia are as large as the pleural and are joined together by a long commis- sure. Two nerves and the cerebral-buccal connective have been observed emerging from each one. The circulatory system consists of a large heart and a single blood gland situated behind the central nervous system (Figure 5A). The renal sac opens directly on the renal pore (Figure 5A). We did not observe glands in the dorsal pericardial walls or in the distal ureter, as Wagele (1989) described for B. hodgsoni and B. clavigera, re- spectively, but that could be due to the poor preservation of our specimens. DISCUSSION The study of additional specimens of Bathydoris aioca in the present paper allows a more detailed comparison with other known species of the genus. The specimens from Oregon show no external or anatomical differences with the holotype from Baja California, and are obviously con- specific. A. Valdés & H. Bertsch, 2000 Page 177 Figure 6 Bathydoris aioca, central nervous system (CASIZ 113315), scale bar = 1 mm. Abbreviations: b, buccal nerve; bg, buccal ganglion; c, cerebral nerve; cg, cerebral ganglion; p, pedal nerve; pc, pedal commissure; pg, pedal ganglion; pl, pleural nerve; plg, pleural ganglion; ppc, parapedal commissure; r, rhinophoral nerve; rg, rhinophoral ganglion. Wagele (1989) reviewed the Antarctic species of the genus Bathydoris Bergh, 1884, comparing them with oth- er species of the genus. She concluded that Bathydoris clavigera Thiele, 1912 (synonyms: B. obliquata Odhner, 1934, and B. argentina Kaiser, 1980), B. hodgsoni Eliot, 1907 (synonyms: B. inflata Eliot, 1907, and B. brownii Evans, 1914), B. vitjazi Minichev, 1969, B. abyssorum Bergh, 1884, B. ingolfiana Bergh, 1900, B. aioca Marcus & Marcus, 1962, and B. patagonica Kaiser, 1980, can be considered as valid species. This conclusion was based on the re-examination of the holotypes of several species and a review of the literature. Only two species, B. cla- vigera and B. hodgsoni were anatomically studied. The holotype of B. aioca was not studied for that paper. In the meanwhile, another new species, B. violacea Bara- nets, 1993, was introduced from Antarctica (Baranets, 1993). Bathydoris aioca is easily distinguished from B. cla- vigera and B. hodgsoni, the two species fully described by Wagele (1989). The radular rachidian teeth of B. cla- vigera have two cusps, which are absent in B. aioca, and the inner lateral tooth is very different in shape from the rest of the laterals, whereas in B. aioca it is very similar. The vaginal duct of B. clavigera is very short and wide, and runs externally to the female gland, whereas in B. aioca it is long and thinner, and runs inside of the female gland mass. Externally, the gill of B. clavigera is in a central position and the body is flattened, whereas the gill of B. aioca is close to the posterior end of the notum, and the body is very high. Bathydoris hodgsoni is also Page 178 different from B. aioca in the radular morphology. The rachidian teeth of the former are triangular and have one cusp, whereas in B. aioca the rachidian teeth are rectan- gular and smooth. Bathydoris hodgsoni shares with B. aioca a long vaginal duct running inside the female gland, the gill situated close to the edge of the notum, and the absence of eyes. Other differences are that the penis of B. hodgsoni has numerous pits and folds, but it is smooth in B. aioca; and the blood gland of B. hodgsoni is situated behind the intestine, whereas in B. aioca it is situated below. Other Antarctic or southern species are B. vitjazi, B. patagonica, and B. violacea, all of them incompletely de- scribed. Bathydoris vitjazi also lacks eyes, but the foot is very small compared to the body. According to Wagele (1989) the number of branchial leaves of B. vitjazi can be interpreted as seven. These features are very different from that of B. aioca, which has a foot as large as the notum and 18 branchial leaves. Bathydoris patagonica is also clearly different from B. aioca. The former has a very large buccal area (see Kaiser, 1980), the body is more flattened than in B. aioca, the foot is larger, the jaws are broader, and the rachidian teeth of the radula are tri- angular, being rectangular in B. aioca. According to Wa- gele (1989), B. patagonica could constitute a geographi- cal variety of B. hodgsoni. Bathydoris violacea is also an eyeless species that differs from B. aioca in the rhino- phore position, which is far anterior in B. aioca and far- ther back in B. violacea (see Baranets, 1993). The radular morphology of both species is also different. The rachi- dian teeth of B. violacea have one cusp, whereas those of B. aioca are smooth. The innermost lateral teeth of B. violacea are very different from the rest, they are wide and short, with a small cusp, whereas in B. aioca the innermost lateral teeth are similar to the other laterals, having an elongate cusp. In addition, the body of B. vio- lacea is elongated, having a posterior prolongation resem- bling a tail. In B. aioca the body is oval and elevated, lacking any posterior prolongation in all specimens ex- amined. The two other species of Bathydoris described from the The Veliger, Vol. 43, No. 2 Pacific Ocean are B. abyssorum and B. ingolfiana. Both differ from B. aioca in details of the external morphology and anatomy. According to Wagele (1989), B. abyssorum has only five branchial leaves (in a 120 mm long speci- men), a small foot, and appears to have two vesicles in the reproductive system; B. ingolfiana has seven bran- chial leaves (in a 90 mm long specimen) and also a small foot, whereas B. aioca has 18 branchial leaves in a 71 mm preserved length specimen, the foot is as large as the notum and has only one genital vesicle. Detailed anatom- ical studies on additional specimens of B. abyssorum and B. ingolfiana are necessary for completion of the diag- nostic features of both species. We were unable to find the visceral loop in the three specimens of B. aioca examined, as well as Wagele (1989), in B. clavigera and B. hodgsoni. According to Wagele (1989), it is probable that it lies very close to the pedal and parapedal commissure within a common sheath of connective tissue. Acknowledgments. This paper has been supported in part by the Ministerio de Educacion y Cultura of Spain (SEUD, through its postdoctoral fellowships program. We are very grateful to our colleague Dr. Igor Subbotin (National University, Los Angeles, California), for translating Baranet’s Russian article for us, and to Dr. Heike Wagele (Ruhr-Universitat, Bochum, Germany) for her comments on the manuscript. LITERATURE CITED BARANETS, O. N. 1993. The primitive nudibranchiate mollusc: Bathydoris violacea sp. n. (Gastropoda, Nudibranchia). Vest- nik Sankt-Peterburgskogo Universiteta (3) 3:13—18 [in Rus- sian]. KAISER, P. 1980. Die Gattung Bathydoris Bergh, 1884 in pata- gonischen Gewiassern. Spixiana 3:43—51. LANCE, J. R. 1967. The holotype of the abyssal dorid nudibranch Bathydoris aoica Marcus & Marcus, 1962. The Veliger 9: 410. Marcus, Ev. & ER. Marcus. 1962. A new species of the Gnath- odoridacea. Anais da Academia Brasileira de Ciencias 34: 269-275. WAGELE, H. 1989. A revision of the Antarctic species of Bath- ydoris Bergh, 1884 and comparison with other known bath- ydorids (Opisthobranchia, Nudibranchia). Journal of Mol- luscan Studies 55:343—364. The Veliger 43(2):179-189 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 Two New Species of the Family Hydrobiidae (Mollusca: Caenogastropoda) from Austria MARTIN HAASE Institut fiir Natur-, Landschafts- und Umweltschutz, Universitat Basel, St. Johanns-Vorstadt 10, CH-4056 Basel, Switzerland ERICH WEIGAND Glatschach 16, A-9772 Dellach/Drau, Austria AND HARALD HASEKE Nationalpark O.6. Kalkalpen Gesellschaft m.b.H. Molln, A-4591 Molln, Austria Abstract. Two new species of the family Hydrobiidae from Austria are described. One belongs to the genus Bel- grandiella A. J. Wagner, 1928, the second is allocated to Bythiospeum Bourguignat, 1882. The crenobiontic Belgrandiella species is characterized by one autapomorphic character state of the genital system and by a unique combination of states which are shared with other species. The second species, a stygobiont, is tentatively attributed to Bythiospeum, because its description is based only on shell characters. Both species are known from very restricted ranges correspond- ing to the distributional pattern of their congeners. In addition, data of 13 abiotic parameters measured over a period of more than 5 years in the springs where the new species were found are presented. The values of these parameters are characteristic for natural, carbonate brooks. Con- tamination with fecal bacteria suggests influence through agriculture or may be caused by feces of mammals. Chemically, agriculture and forestry appear to have no significant impact on the springs investigated. INTRODUCTION Throughout the world, karst systems are known to be in- habited by speciose faunas including many stygobionts (e.g., Ilhes, 1978; Botosaneanu, 1986; Juberthie & Decu, 1994). Since 1991, the beginning of the karst monitoring projects of the area of the then prospective, now estab- lished (25 July 1997) Nationalpark Kalkalpen and its sur- roundings in the Northern Calcareous Alps of southeast- ern Upper Austria (Atzwanger, 1993; Haseke, 1993), a number of hydrobioid (sensu Davis, 1979) gastropods were found. The species belong to the genera Belgran- diella A. J. Wagner, 1928, and Bythinella Moquin-Tan- don, 1855, both largely crenobiontic genera, and the sty- gobiontic genera Bythiospeum Bourguignat, 1882, and Hauffenia Pollonera, 1898. The Belgrandiella and By- thiospeum species, one species each, turned out to be new to science. The monitoring not only comprises faunistic recordings but also the collection of physical parameters of the springs. Thus, we are able to present detailed descriptions of the environment of the new hydrobiids, which is an important step toward understanding the autecology of crenobiontic and stygobiontic organisms. MATERIALS anpD METHODS All four localities where the snails presented in this study have been collected are in the drainage area of the rivers Steyr or Krumme Steyrling, a left tributary of the Steyr, which empties into the Enns shortly before the latter flows into the Danube. This river system is part of the right (southern, Alpine) catchment area of the Danube. More detailed locality data and information about the to- tal number and the number of specimens investigated are given in the systematic descriptions. Physical parameters of the springs measured are listed in Table 3. These data were taken from Haseke (1998). The material is deposited in the Museum of Natural History in Vienna (NHMW). The snails were fixed in Bouin’s solution, in 4% form- aldehyde or in 70% ethanol, respectively. Anatomical methods were those of Haase (1992). The SEM pictures were partly made on a JEOL JSM 35 Scanning Micro- scope at the Zoological Institute in Vienna and partly on Page 180 a JSM 6300 at the SEM Laboratory of the University of Basel. SYSTEMATIC DESCRIPTIONS General Remarks In the present descriptions of the new species, only characters relevant for the diagnosis of species are pre- sented. The descriptions are based on fixed material. Therefore, several characters whose investigation requires observation of living animals could not be included (cf. Hershler & Ponder, 1998). Whenever possible, we used the terminology of Hershler & Ponder (1998) for the def- inition of character states. Only the material from the type localities is declared type material in order to avoid tax- onomic confusion in case the populations from other lo- calities should turn out not to be conspecific, applying finer methods like molecular techniques. Belgrandiella A. J. Wagner, 1928 Type species: Belgrandia kusceri A. J. Wagner, 1914 Diagnosis: Shell pupiform, rarely turriform, usually smaller than 2.5 mm; protoconch pitted, teleoconch with- out sculpture. Stomach without caecum. In females the intestine lies close to the pallial oviduct with only a weak- ly developed loop to the left; in males this loop is quite distinct. Female genital system characterized by a simple ovary, the presence of a bursa copulatrix, and a recepta- culum seminis lying ventral to the bursa; renal oviduct describes a single, wide loop before passing into the pal- lial oviduct. In males the testis has simple, wide lobes; vas deferens enters and leaves the prostate at its very ends; ejaculatory duct only weakly developed; penis without glands but often a muscular lobe on the left side. Remarks: This diagnosis is based on Radoman (1983) and Haase (1994). The entire anatomy of a Belgrandiella species is presented in Haase (1993a). However, the de- limitation of Belgrandiella from other, similar nominal genera is ambiguous, and concepts of various authors dif- fer considerably. Lack of knowledge of anatomical data of many species attributed to various nominal genera and nomenclatural problems have prevented a thorough sys- tematic analysis and clear classification of this group of species so far. This problem is outlined in more detail by Haase (1996) in his discussion of the generic allocation of the Austrian radiation attributed to Belgrandiella. Belgrandiella aulaei Haase, Weigand & Haseke, sp. nov. (Figures 3—7) Holotype: NHMW 89958. Paratypes: NHMW 89959 (one series of histological sec- tions, >50 specimens). The Veliger, Vol. 43, No. 2 Type locality: Rinnende Mauer, a system of springs emerging on the foot of a slightly overhanging wall of conglomerate overgrown with mosses respectively on top of that wall over a length of about 30 m forming a beau- tiful curtain of drops (Figures 1, 2). This system of springs is close to the left bank of the river Steyr north of Leonstein 405 m above sea level. The snails live in the lower springs and their common discharge. Additional material: Right spring of the Wunderlucke (Figure 1), a pond, close to the left bank of the river Krumme Steyrling in Rabach, 365 m above sea level (NHMW 89960, four series of sections, >25 specimens). Etymology: Au/aeum (Latin, neuter) means splendid cur- tain and refers to the curtain of drops at the type locality. Diagnosis: B. aulaei is characterized by the position of the receptaculum seminis between albumen gland and bursa copulatrix and by the unique combination of the following characters: capsule gland bipartite, posterior al- bumen gland asymmetrical, bursa copulatrix behind al- bumen gland, bursal duct as wide loop. Description: Shell. Pupiform, shallow sutures, transpar- ent, with up to 3.75 moderately convex whorls of which the pitted protoconch comprises about one whorl (Figures 3, 4). Aperture obliquely ovoid; outer lip straight and or- thocline. Umbilicus a slit. Measurements are given in Ta- ble 1. Operculum. Thin, orange, paucispiral, nucleus submar- ginal. Radula (Figure 5): R: 5-7 1 5-7/1 1 L: 3-4 1 5-6 M;: 24-26 M,: 24-29. Central tooth trapezoidal; basal tongue of the central tooth as long as lateral margins and V- shaped, with curved edge; basal cusps prominent. Face of lateral teeth taller than wide, with basal projection; lateral wing much longer than cutting edge. Cusps on inner marginal teeth slightly larger than on outer marginal teeth; cutting edge on inner marginal teeth longer than 25% of total tooth length. Non-genital anatomy. In most specimens mantle en- tirely black. But the epithelium lying over the distal glands of the genital system (prostate, pallial oviduct) and the stomach may have less pigment. Epidermis of head and foot practically unpigmented. But their connective tissues as well as the radular bolsters and the pharynx contain black granules. Cephalic tentacles bear no cilia. In one female from the Wunderlucke two additional eyes were found on the base of the right tentacle. No ctenid- ium. Hypobranchial gland very short lying in the rear of the mantle cavity. Endothelium of the stomach may con- tain some granules of black pigment. The stomach and intestine of one individual from the Wunderlucke were full with diatoms. Male reproductive system. Base of the penis broadened on the left side. A round to elongate lobe is more or less well separated from this base (Figure 6). Behind the tip, M. Haase et al., 2000 Page 181 ae. GS Gee 372 Wendsqen—Bahnhofsdlg Stiedelsbach —Pechgrabe | uf aN 1273 Kienberg 3 a RIM a Hohe, Dirn Diirnberg 1208, as ia Eunen C a 1285 1134 Schieferstein radau Set | A hoberstein A Leonstein " ep /WULU Se wy Ks Arzberg Ss 7 A ne” ae. Arn rierl ie ee jonrbe’ 358 SS Heinz waldstein Pe eee ee Be ss ne ee PG Blumen / Reichraming g iC Ree chet f- -leonstein Molln Reilern + orn Sulzbach Dirnbach CEI KAEE OU Carey Breitenau ~ Hedelsbach 2 Oberésterreic: Zimeck ) 1244 a \ Schneeberg e Unt.- S Garnweid = ae \ -Wienerweg Eibling ES Weibenbach g gy on ae = Hoher Traiing Hausbach Ss - Schon & 1108 Ale & oe f b Wee Frauenster hlapfengraben S Sn ee ae WEL (anzennech s cf Dirngraben = : bauer jf Tera \, Effertsbach IL : ‘ runnbac = R auer Grdftenbg. : : : SS je x1990 S ‘aie ee Steyern | 4. Epedigtsiny! 509 4 \ 2. d. Pyhrnbahn Gr.Spitzberg a Sy Be. Zobel 4 } ! ott pee Wo ~. es Be % \a 1305 N ro (“ / Klauser See = ( eas REUT A ‘ f ‘ : ee aN / tay es r= og mele A pee et x = : . : é : | g re 1580 _Aipstein 8 : 4838 ¢ : 5 : : 3433 ee te | Hochsen a g _ Bodinggraben\, | a os he : Fee C8 os : oss ee i ee Hochkogel ae Ne : : : LS 6 — CoN 1157 x \ SER Hoher Nock / : : ee : - . 2S Dirnbach ee : 02h Be Brandriegel Gamspien Gf = © sykbriicke 1474 OUR RR ce SATA ns nets saat 7.3. The Rinnende Mauer and the Reutersteinquelle are well oxygenated with relative concentrations of over 90%. The spring Welchau, which may even be oversat- urated with oxygen as the preceding two springs, shows quite large fluctuations in oxygen concentration, whereas Figure 8 Shells of Bythiospeum nocki Haase, Weigand & Haseke, sp. nov. from the Reutersteinquelle. A, Holotype (NHMW 89961); B—-E, Paratypes (NHMW 89962). Shell height of the holotype = 1.47 mm. Page 186 The Veliger, Vol. 43, No. 2 Table 3 Physical parameters of the springs. Abbreviations: C = electrical conductivity (uS/cm), min = minimum, max = max- imum, N = number of measurements, Q = discharge (L/s), relO, = relative concentration of oxygen (%), T = temper- ature (°C). All concentrations are given in mg/L. Spring, period Q T pH O, relO, Rinnende Mauer N 20 19 18 Vf 6 5/22/92—20/8/97 min 7.0 6.8 7.39 8.90 95.0 max 25.0 10.7 8.25 11.55 102.0 Wunderlucke N 22 22 19 7, 6 7/18/9 1—20/8/97 min 0.5 3.6 7.36 5.30 79.0 max 30.0 13.7 8.26 9.80 89.0 Reutersteinquelle N 20 10 9 5 5 7/18/91—15/5/97 min 0) 6.1 7.53 10.70 92.0 max 200.0 6.7 8.30 13.50 116.0 Welchau N 19 18 17 6 5) 7/18/9 1—20/8/97 min 0.0 6.8 7.35 7.50 66.0 max 295.0 8.0 8.15 11.62 102.4 G | Mp= © x24 (Nav ke NOs SO Sole 19 20 20 15 15 20 18 17 ME NO ONS OA) WO G67 O55 343. 31.96 70107 1.14 “0189 1A07m IS tOommEIG a I), 7D 22 11 1 22 18 16 348 (0102 38:77 0.25 (003). SDS ous 571 22.97 126.0 HO OL 7 SATA O73 10 10 10 5 5 10 7 7 213. 656 2737 0.15. O18) S:O7 eS OOMmNO NID 265 13.19 4330 034 004) eyOmE OSD 18. 18 18 9 9 18 16 14 313 15.85 3278 0.19.) (028) oa Sum OO 380', 23.60) 51:00) | 0:73) 0.42 eemlG 0.63 i : : : Ao 15.70 the Wunderlucke fluctuates in a quite small range but nev- er reaches saturation. Ion concentrations and hence elec- trical conductivity vary most strongly in the Wunder- lucke. Especially remarkable are the minimum concentra- tion of 0.02 mg/L of magnesium and the peak at 84.74 mg/L of sulfate. DISCUSSION The Austrian species of both genera Belgrandiella and Bythiospeum have been the subject of recent revisions (Haase, 1995, 1996). The finding of the new Belgran- diella species extends the known range of this genus to- ward the west, and brings the range of the extant species closer to the extinct B. intermedia Boeters (1970) from the Tiefsteinschlucht near Salzburg, whose generic allo- cation is presumptive due to the lack of anatomical data (cf. Haase, 1996). In their distributional pattern both new species correspond to their congeners (Haase, 1995, 1996) in that their ranges are very narrow being, as far as we know today, restricted to two springs and the drain- age area of the Krumme Steyrling. The type locality of B. aulaei, Rinnende Mauer, has been threatened through a gravel mining project and plans to build an incinerating plant and a waste disposal site in its drainage area. Protests of the local population have stopped these projects, at least for the time being. The company involved has already announced plans to seek approval of a modified project (Maier & Maier, 1997). We hope that the discovery of the new species will strengthen arguments for protection of this area, which has been the aim of local organizations since 1983 (Maier & Maier, 1997). Furthermore, the finding of two new, presumably highly endemic species in the surroundings of the Nationalpark Kalkalpen, demonstrates that also this surrounding area is faunistically peculiar, suggesting an extension of the borders of the national park. To those species ascribed to Belgrandiella tentatively because of lack of anatomical data listed in Haase (1996), viz. B. intermedia and B. styriaca Stojaspal, 1978, two more species, B. multiformis Fischer & Reischtitz, 1995, and B. kreisslorum Reischitz, 1997, both from the south- east of Austria, were added recently (Fischer & Reis- chiitz, 1995; Reischiitz, 1997). But these species are not identical to those represented by samples deposited in the NHMW mentioned in Haase (1996). These two unde- scribed species, as well as that mentioned in Haase (1993a), have still not been rediscovered in the field and therefore we refrained from a formal description. Wheth- er the fossil B. dehmi Boeters, 1995, from Oberfranken in Germany is a true Belgrandiella will remain subject to speculation, although its shell shape and occurrence sug- gest a close relationship to the Austrian species (Boeters, 1995). In general, very little is known about the biology of freshwater hydrobiids in contrast to their brackish water relatives of the genus Hydrobia Hartmann, 1821 (e.g., Fretter & Graham, 1978; Lassen & Clark, 1979; Davis et al., 1989; and literature cited therein), and Potamopyrgus antipodarum Gray, 1843 (e.g., Duncan & Klekowski, 1967; Winterbourn, 1969; Ponder, 1988; Jacobsen & Forbes, 1997; Jokela et al., 1997; and literature cited therein), which tolerates a wide range of salinities from brackish to freshwater. Bythinella dunkeri (Frauenfeld, 1856) is the only species of hydrobioid spring snails in which feeding was systematically investigated (Oswald et al., 1991; Brendelberger, 1992, 1995). Diatoms were among the preferred food sources beside cyanobacteria, bacteria, hyphomycetes, and certain algae. Diatoms ap- pear to be an important component of the diet also of Belgrandiella aulaei as may be judged by the contents of the stomach and intestine of one specimen. The values of the 13 parameters measured in all four M. Haase et al., 2000 springs are characteristic for natural carbonate brooks and groundwaters (e.g., Drever, 1982; Otto & Braukmann, 1983; Braukmann, 1987; Kummert & Stumm, 1989). However, in all springs, fecal bacteria such as Escherichia coli, other coliform bacteria, and enterococci were de- tected (Schmidt, personal communication). This contam- ination may be due to agricultural influences in the drain- age areas of the Rinnende Mauer and the Wunderlucke, or, in case of the Reutersteinquelle and the spring Wel- chau, caused by feces of deer and other mammals. Chem- ically, agriculture and forestry appear to have no signif- icant impact on the springs investigated, at least not on the parameters measured. The relatively high fluctuations of the various concentrations measured in the Wunder- lucke may be due to the influence of the nearby Krumme Steyrling at highwater. That holds also for the occasion- ally low concentrations of magnesium. The high peaks of sulfate in this spring may indicate that part of the water drains from rock strata containing gypsum (Tollmann, 1985) at times. Because of the extreme fluctuations of the discharge ranging from 0 to 200 and more L/sec, crenobiontic spe- cies cannot establish permanent populations in the Reu- tersteinquelle and the spring Welchau. In the Reuterstein- quelle only a second stygobiontic species of the genus Hauffenia was found. Hauffenia sp. was also washed out of its habitat in the spring Welchau. Nevertheless, some specimens of a Bythinella species were also found here probably indicating recent recolonization from a nearby source population. At the Rinnende Mauer and in the Wunderlucke, both with moderate and permanent dis- charge, B. aulaei was accompanied by a Bythinella. Since there is, to our knowledge, no comparable study as to the number of physical parameters measured in a spring inhabited by hydrobioids and as to the continuity of the data collection over a period of more than 5 years, we are not able to bring these data into a wider context of general ecology of crenobiontic hydrobiids. Occasion- ally, temperature data, pH values, and/or electrical con- ductivity are given (e.g., Bregenzer, 1915; Girod & Pez- zoli, 1967; Boeters, 1969, 1970, 1977; Jungbluth, 1972; Ponder, 1989; Hershler, 1998). The majority of those spe- cies for which this kind of information is available live in similarly cool waters as the two species dealt with in this paper. However, a number of species inhabit con- stantly warm or even thermal springs with temperatures of 19°C and more (e.g., Neohoratia ateni [Boeters, 1969]; 33°C [Boeters, 1969]; Belgrandiella parreyssii |L. Pfeif- fer, 1841]: 23.5°C [Boeters, 1970]; B. mimula: 19.5°C [Boeters, 1970], cf. Haase [1996] for the identity of this species]; Heleobia aponensis (Martens, 1858]: ca. 30°C [Boeters, 1977]; several unnamed Australian species: up to 46°C [Ponder, 1989]; several species of Pyrgulopsis Call & Pilsbry, 1886: up to 36°C [Hershler, 1998]). Os- wald et al. (1991) observed that the hydrobioid Bythinella dunkeri reproduced also at room temperature although the Page 187 original habitat had a mean temperature of only 8.8°C. This finding contrasts with frequent statements that By- thinella species are cold stenothermic (e.g., Bregenzer, 1915; Jungbluth, 1972) and suggests that low tempera- tures might not necessarily be a prerequisite for hydro- bioid crenobionts for the colonization of a spring. We rather assume that the current of a spring may not exceed a certain velocity so that both the food sources of the snails and the snails can establish populations and are not swept away. In addition, competition and predation pres- sure may also play an important role for the restriction of certain hydrobiids to the crenal region of a brook (Os- wald et al., 1991). The only stygobiont hydrobiid species we are aware of for which measurements of temperature and oxygen con- centration were made directly in their habitat, i.e., in the groundwater, are Lobaunia danubialis Haase, 1993, and Bythiospeum cf. geyeri from the groundwaters of the river Danube in Vienna (Pospisil, 1989, who referred to these species as Horatia sp. and Paladilhiopsis geyeri, respec- tively; as to the identity of these species see Haase, 1993b, 1995). The temperatures of 8.6 to 12.4°C in the Danube groundwater are well in the range measured for the majority of European crenobiontic hydrobioids. But the oxygen concentrations of 0.042 to 0.7 mg/L are re- markably low and in strong contrast to the oxygen situ- ation in springs. It can be assumed that the oxygen con- centration in the real habitats of B. nocki is much lower than measured in the springs where it has been found, while the measurements of the other parameters may well reflect the actual conditions in the groundwaters draining to the Krumme Steyrling. Acknowledgments. We are grateful to the following people from the Nationalpark 0.6. Kalkalpen Gesellschaft m.b.H. Molln for their cooperation: Erich Mayrhofer, Elmar Proll, Roswitha Schrutka, and Norbert Steinwendner. This Gesellschaft also gave permission to publish data from internal reports. Susanne Schmidt (Graz) shared her bacteriological data with us. Klement Tockner’s (now Ziirich) help in the field is acknowledged. We thank Richard Guggenheim and Waltraud Klepal, heads of the SEM laboratories of the Universities of Basel and Vienna, re- spectively, and their staff for providing the facilities for SEM and for their help. Thanks also to Daniel Wey (Basel), who did the darkroom work. Financial support was received from the Na- tionalpark O.6. Kalkalpen Gesellschaft m.b.H. Molln. LITERATURE CITED ATZWANGER, M. 1993. Geplanter Nationalpark Kalkalpen. Un- terschutzstellung und vernetztes Forschungsprogramm. Wis- senschaftliche Beihefte zur Zeitschrift Die Hohle 42:109— 116. BERNASCONI, R. 1990. Revision of the Genus Bythiospeum (Mol- lusca Prosobranchia Hydrobiidae) of France, Switzerland and Germany. Privately published: Miinchenbuchsee. 44 pp., 19 figs. Boeters, H. D. 1969. Pseudamnicola klemmi n. sp. aus Stid- frankreich und Microna ateni n. sp. aus den spanischen Pyr- Page 188 The Veliger, Vol. 43, No. 2 enden (Mollusca, Prosobranchia). Archiv fiir Molluskenkun- de 99:69-72. Boeters, H. D. 1970. Die Gattung Microna Clessin, 1890 (Pro- sobranchia, Hydrobiidae). Archiv ftir Molluskenkunde 100: 13-145. Boeters, H. D. 1977. Westeuropadische Hydrobiidae, 6. Hydrobia (Semisalsa) Radoman (Prosobranchia). Archiv ftir Mollus- kenkunde 108:45—50. Boeters, H. D. 1995. 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Rio San Pedro s/n, Apdo. 40, 11510 Puerto Real (Cadiz), Spain JOSE CARLOS GARCIA-GOMEZ Laboratorio de Biologia Marina, Departamento de Fisiologia y Biologia Animal, Facultad de Biologia, Universidad de Sevilla, Av. Reina Mercedes 7, Apdo. 1095, 41080 Sevilla, Spain AND RICCARDO CATTANEO-VIETTI Istituto di Zoologia, Universita di Genova, via Balbi 5, 16126 Genova, Italy Abstract. Useful additional data on the dorid nudibranch Tambja simplex Ortea & Moro, 1998, are given. The external and internal features of this species are compared with those of the single specimen of the original description and those of the most similar Atlantic species, 7. marbellensis Schick & Cervera, 1998. Tambja simplex is blackish purple with a yellow edge on the notum and the foot, and only very few bands of differing length of the same color on the notum, flanks, and tail. There is also yellow coloration on the inner and outer sides of the rachis of the gills and on the upper edge of the oral tentacles. The radula is typical of the genus Tambja, with the inner lateral tooth having a conspicuous denticle on the inner edge of the primary cusp. The second inner lateral tooth also has a small cusp. The reproductive system has a prostate differentiated from the deferent duct, a rounded bursa copulatrix, and an elongate seminal receptacle, smaller than the bursa copulatrix. INTRODUCTION To date, the knowledge of the opisthobranch fauna of the Cape Verde Archipelago is limited. Recently, Cervera et al. (1996) gave an account of all existing references re- ferring to the opisthobranchs from this archipelago, to which we have to add Ortea & Pérez (1992), Martinez et al. (1996), Ortea et al. (1996, 1997). Moreover, Schick & Cervera (1998) gave an account of the known Atlantic species of the genus Tambja Burn, 1962. This included the only two known species from the coasts of the Cape Verde Archipelago to date, 7. fantasmalis Ortea & Garcia-Gomez, 1986 and T. anayana Ortea, 1989. During a brief survey of Uha do Sal, Cape Verde Islands in De- cember 1990 and January 1991, one specimen belonging to an undescribed species of this genus was collected by one of us (R.C.V.). Another specimen of the same species was collected in July 1997 during the sampling of the project ‘“‘Evaluacion de los recursos naturales litorales de la Republica de Cabo Verde,”’ carried out by the Depart- ment of Biology of the University of Las Palamas de Gran Canaria (Spain). However, while a full description of this species was in press, two papers with more infor- mation about the opisthobranchs from Cape Verde Islands were published (Ortea & Espinosa, 1998; Ortea & Moro, 1998). This last paper included, on the basis of a single specimen, the description of our undescribed Tambja that we had in press. The present paper provides useful ad- ditional information on T. simplex Ortea & Moro, 1998, including six more specimens recently collected at the same archipelago by Dr. Peter Wirtz (University of Ma- deira, Portugal). SYSTEMATIC DESCRIPTION Suborder DORIDACEA Family POLYCERATIDAE Alder & Hancock, 1845 Genus Tambja Burn, 1962 Tambja simplex Ortea & Moro, 1998 (Figures 1-3) Material: One specimen, 60 mm in length, collected at 15 m depth, Porto do Anciao, Ilha do Brava, Cape Verde Archipelago, eastern Atlantic, July 1997. This specimen has been deposited at the California Academy of Sciences (San Francisco, USA), with the catalogue number CASIZ 110368. One specimen, 40 mm in length, collected under a stone at 30 m depth, Albacora Bay, Ilha do Sal, Cape J. L. Cervera et al., 2000 Page 191 ye Figure 1 Tambja simplex. Dorsal view (A), ventral view (B), and lateral scheme (C) of specimen MNCN 15.05/29571. D. Inner view of a gill. E. Color pattern of the outer side of the rachis of the gills. Key: dpu, dark purple; ye, yellow. Figures D and E based on more than one specimen. Verde Archipelago, eastern Atlantic, January 1991. This specimen has been deposited at the Museu Nacional de Ciencias Naturales (Madrid, Spain), with the catalogue number 15.05/29571. Six specimens, 11-30 mm in length (preserved), col- lected at 10—35 m depth on bryozoans, Tarrafal, Ilha do Sao Tiago, Cape Verde Archipelago, December 1998. These specimens have been deposited at the Museo Na- cional de Ciencias Naturales (Madrid, Spain), with the catalogue number 15.05/33320. Diagnosis: Body limaciform with widened head; notum with smooth edge and tail. Ground color black-purple. Edge of notum and foot yellow; notum with a middle yellow line; back of the tail yellow; flanks with two yel- low lines, the wider of which joins anteriorly and poste- riorly with that edging the foot, and then extends on upper edge of oral tentacles. Outer surface of tip of rachis of gills yellow and yellow line also present on inner surface of rachis of these structures. Rhinophoral sheaths bor- dered with yellow. Rachidian tooth wider than tall and notched at anterior edge; inner lateral radular tooth hooked, with large bicuspid primary cusp; second inner lateral tooth with small cusp. Bursa copulatrix and sem- inal receptacle different in size; prostate well differenti- ated and vestibular gland very large. Description: Body limaciform with widened head; notum and its edge and tail smooth. Oral tentacles short and dorsoventrally flattened. Rhinophores with 18—19 lamel- lae. Five gills tripinnate and non-retractile, surrounding anal papilla. Three anterior gills more highly developed. Page 192 The Veliger, Vol. 43, No. 2 5mm Imm Figure 2 Tambja simplex. A. Buccal mass. B. Labial cuticle. C. General view of the radula. D. Detail of the lateral and marginal radular teeth. E. Detail of a dorsal view of the primary cusp of the lateral radular teeth. Figures A and B based on more than one specimen; figures C, D, and E based on MNCN 15.05/29571. Key: oe, esophagus; sgl, salivary glands. Ground color black-purple. Notal edge marked by thick yellow band, interrupted posteriorly, but continuing along back of tail to its end. Broken stripe of same color run- ning from bases of rhinophores toward gill tuft (Figure 1A). Edge of foot bordered by yellow band (Figure 1B). On each flank of body, between notal edge and that of foot, two other lines of same color present, upper broken and thinner than lower, which is uninterrupted. Latter joining yellow band of edge of foot anteriorly and pos- teriorly; then, both extending to upper edge of oral ten- tacles (Figure 1B, C). Rhinophores black-purple; rhino- phoral orifices surrounded by yellow ring. Gills black- purple, but inner side of the rachis of each one having yellow line connected between them (Figure 1A, D). Large yellow patch on outer upper side of each gill rachis, which may reach almost their bases (Figure 1E). Internal anatomy. Salivary glands on buccal mass flank- ing esophagus (Figure 2A). Labial cuticle becoming stronger just at buccal aperture region (Figure 2B). Rad- ular formula of Porto do Anciao specimen 17 X 4.1.R.1.4 and that of Albacora Bay specimen (Figure 2C) 20 X 4- J. L. Cervera et al., 2000 ps hd sr be vg! Figure 3 Tambja simplex. Reproductive system. Key: am, ampulla; be, bursa copulatrix; dd, deferent duct; fgl, female gland; hd, her- maphroditic duct; p, penis; pr, prostate; ps, penial sheath; sr, sem- inal receptacle; va, vagina; vgl, vestibular gland. Figure based on CASIZ 110368 and MNCN 15.05/29571. 3.1.R.1.3-4. Rachidian tooth much wider than tall, notched at anterior edge. Inner lateral tooth hooked with large primary cusp, with strong denticle on its inner edge, and smaller triangular basal cusp (Figure 2D, E). Outer lateral teeth scalelike and less developed than the former, although second inner lateral tooth with small cusp (Fig- ure 2E). Reproductive system (Figure 3) with similar ar- rangement in both dissected specimens, with hermaph- roditic duct that continues as S-shaped ampulla. Bursa copulatrix rounded; seminal receptacle, which is smaller than bursa, elongate. Oval vestibular gland very well de- veloped and much larger than two above sacs. Deferent duct with well differentiated prostate. Penis armed with numerous hooked spines. DISCUSSION Recently, Schick & Cervera (1998) presented a compar- ative table of the Atlantic species of Tambja. In the In- troduction, we have given the history of the description of this species. The external anatomy, coloration and rad- ula described by Ortea & Moro (1998) are in agreement with those of our specimens. The reproductive system is described for the first time in this paper. The ground color and the presence of yellow stripes on the notum and flanks and the yellow edge of the notum of 7. simplex permit us to separate it from 7. gratiosa (Bergh, 1890), T. capensis (Bergh, 1907), T. divae (Marcus, 1958), T. oliva Meyer, 1977, T. fantasmalis Ortea & Garcia-Go- mez, 1986 and, T. anayana Ortea, 1989. Tambja ceutae Garcia-Gomez & Ortea, 1988, also has yellow stripes on Page 193 the notum and the flanks, but in a higher number than 7. simplex and always bordered with dark blue-black. More- over, T. ceutae has conspicuous blue or greenish blue conical papillae on the edge of the notum and the tail. The most similar species to T. simplex is T. marbellensis Schick & Cervera, 1998, but this latter species has fewer yellow lines on the notum and the flanks than 7. simplex. Additionally, in the adult specimens of 7. marbellensis all the yellow marks are shaded by brown, and those that are on the outer side of the rachis of the gills are usually more spread than in 7. simplex, never reaching the top of the gill. The gills of the latter species lack yellow col- oration on the inner surface of each rachis and lack yel- low marks on the upper edge of the oral tentacles. With respect to the internal anatomy, the rachidian radular tooth in T. simplex, as stated by Ortea & Moro (1998), is much wider than tall, whereas in 7. marbellensis it is almost as wide as tall. The arrangement of the reproduc- tive system of both species is very similar, but in 7. sim- plex the prostate is less enlarged and the vestibular gland is more developed than in T. marbellensis. Moreover, in T. simplex the seminal receptacle is elongate and smaller than the bursa copulatrix, whereas in 7. marbellensis it is pyriform and similar in size to the bursa copulatrix. The arrangement and shape of the penial spines have not been studied in either species. Schick & Cervera (1998) discussed the exact identity of the species 7. diaphana (Bergh, 1878) described by Provot-Fol (1927) from the Moroccan coast and stated that it could constitute an additional new species of this genus, although it would require the study of additional specimens from the same locality or nearby areas. How- ever, the sulphur yellow ground color with four emerald lines on the notum, that join anteriorly and posteriorly, distinguishes T. simplex from Pruvot-Fol’s species. Acknowledgments. We want to acknowledge Fernando Espino for collecting the Porto do Anciao specimen and providing its photograph, and Dr. Emilio Roldan for giving us data on its ex- ternal features. This specimen was collected during the field sam- pling of the project “‘Evaluacion de los recursos naturales litor- ales de la Republica de Cabo Verde,” supported by the Depart- ment of Territorial Policy and Environment of the Canarian Re- gional Government, within its framework Program of International Cooperation. Our sincere gratitude to Drs. Terrence M. Gosliner and Barry Roth for the critical reading of the man- uscript. We also thank Dr. Peter Wirtz for providing six additional specimens and their photographs, Mr. Agustin Santos for his help during the preparation of the manuscript, and Mr. José Maria Geraldia from the Electron Microscopy Service of the University of Cadiz for providing facilities to take the Scanning Electron Microscope photographs. LITERATURE CITED CERVERA, J. L., R. CATTANEO-VIETTI & M. EDMUNDs. 1996. A new species of notaspidean of the genus Pleurobranchus Cuvier, 1804 (Gastropoda, Opisthobranchia) from the Cape Verde Archipelago. Bulletin of Marine Science 59(1):150— SVs Page 194 MartTINEZ, E., J. ORTEA & M. BALLESTEROS. 1996. Redescription of Geitodoris reticulata Eliot, 1906 from the Cape Verde Islands. Journal of Molluscan Studies 62:257-261. OrTEA J. A. & J. M. PEREZ. 1992. Captura de Plocamopherus maderae (Lowe, 1842) (Mollusca: Nudibranchiata) en los archipiélagos de Canarias y Cabo Verde. Actas del V Sim- posio Ibérico de Estudios del Bentos Marino 2:229—235. OrtTEA, J. A. & J. Espinosa. 1998. Estudio de nueve especies del] género Flabellina Voigt, 1834 (Mollusca: Nudibranchia) Co- lectadas en Angola, Cabo Verde, Costa Rica, Cuba y Por- tugal, con la descripcion de tres especies nuevas. Avicennia, 8/9:135-148. OrTEA, J. A. & L. Moro. 1998. Descripcion de tres moluscos nuevos de las islas de Cabo Verde. Avicennia, 8/9:149—154. The Veliger, Vol. 43, No. 2 OrTEA, J. A., L. Moro & J. Espinosa. 1997. El género Doto Oken. 1815 (Mollusca: Nudibranchia) en las islas Canarias y de Cabo Verde. Avicennia 6/7:125—136. OrTEA, J.. A. VALDES & J. C. GARC{A-GOMEZ. 1996. Revision de las especies atlanticas de la familia Chromodorididae (Mol- lusca: Nudibranchia) del grupo cromatico azul. Avicennia, suplemento 1:1—165. PRuvot-Fot, A. 1927. Sur quelques mollusques nudibranches de la cote atlantique du Maroc. Bulletin de la Societé des sci- ences naturelles du Maroc 7:39—49. Scuick, K.-L. & J. L. Cervera. 1998. Description of a new spe- cies in the genus Tambja Burn, 1962 (Gastropoda: Nudi- branchia: Polyceratidae) from southern Spain. The Veliger 41(4):344-350. The Veliger 43(2):195—196 (April 3, 2000) THE VELIGER © CMS, Inc., 2000 NOTES, INFORMATION & NEWS The Egg Capsule and Young of the Gastropod Pyrulofusus dexius (Dall) (Buccinidae: Volutopsiinae) Vladimir V. Gulbin Institute of Marine Biology, Vladivostok 690041, Russia Introduction Four species of the genus Pyrulofusus Moerch, 1869, are known currently. These are P. deformis (Reeve, 1847), P. harpa (Moerch, 1857), P. dexius (Dall, 1907), and P. melonis (Dall, 1891) (Kantor, 1990). All of them occur in the North Pacific Ocean. However, egg capsules are known only for P. deformis (Gonor, 1964) and P. harpa (Cowan, 1965). This work describes the egg capsule and young specimens of Pyrulofusus dexius (Dall, 1907). Materials and Methods The material was collected in the northeastern Sea of Okhotsk in August-November 1997 during an explor- atory fishing voyage of the trawler Karatau engaged in the study of buccinid gastropods (collectors V. Gulbin and A. Maltsev). Mollusks were collected from a depth of 180-280 m using special traps of Japanese design. Fish was used as a bait. A total of 400 stations was made. Unlike most Buccinidae, species of the genus Py- rulofusus are predators. They feed on echinoderms, mainly holothurians (Kantor, 1990) and therefore sel- dom get into traps. For this reason, only at three stations (sta. 14, 19, 61) out of 400 were a single species of Pyrulofusus, P. dexius (Dall, 1907) found, and at one station (sta. 6) its egg capsule (Table 1). At these sta- tions, the near-bottom water temperature was 2.8—3.2°C. Earlier (Kantor, 1990), this species was recorded off the coast of Hokkaido, the southern and northern Kurils, the southern coast of Kamchatka, and off the Aleutian Is- lands. This is the first record of this species in the north- ern Sea of Okhotsk. The egg capsule (No 3/46838) is deposited with the gastropod collection at the Institute of Marine Biology (Vladivostok, Russia). Results and Discussion The egg capsule (Figure 1A, B) was found in the north- eastern Sea of Okhotsk near the western coast of Kam- chatka, at a depth 257-265 m on silty sand substratum. The capsule is low, dome-shaped, and firmly cemented onto the surface of a small stone by its broadest surface, the base. At this point, the capsule is quite circular and has a diameter of 58 mm. A circular fringed band of whitish material surrounds the base and is 1-6 mm wide. Height of the capsule is 24 mm. It is composed of con- chiolin. The capsule is yellowish ashen in color, hard, about 0.2—0.25 mm thick, and its surface is clearly marked with a very fine pattern resembling a dense net- work of rectilinear wrinkles 0.3-2 mm long, crossing each other at acute angles (Figure 1C). A coat of lime, covering freshly deposited capsules (Kantor, 1990) was not preserved. Inside, there is a thinner, more flexible smooth inner layer, which is semitransparent and makes visible the outer surface pattern. The capsule base is thin (approx. 0.15 mm), membranelike, semi-transparent, and lacks a pattern. The Pyrulofusus capsule has no special devices for re- leasing the young, such as protein “‘corks,”’ dissolving by the time of maturation in Urosalpinx cinerea (Hyman, 1967), or a cap, opening by hatching time in Trophon (Thorson, 1946). Presumably, the hatchling mollusks can gnaw through the capsule wall at any place. The egg capsule contained three young specimens of near-hatching age. Not a trace remained of any nurse eggs that may have been present. As is known, apart from nurse eggs, there is an additional food source of embry- os—the protein layer of the capsule. This assumption is supported by the fact that the protein layer was absent in the capsule. Three young specimens found in the capsule were pale lilac, with a well-developed, dextral calcareous shell 20— 21 mm high and 13.0—13.5 mm in diameter. They had 2.75 whorls. The first two whorls were almost smooth, except for rare incremental lines and a faint indication of spiral sculpture. The third whorl was distinguished by its pronounced spiral sculpture of 30-32 ridges. In addition, Table 1 Characteristics of stations where the shells and egg capsule Pyrulofusus dexius were found. Coordinates Station no Date (lat. N.—long. E. 6 5 August 1997 57°59'—155°44' 14 7 August 1997 57°48'-155°57' 19 10 August 1997 58°46’-154°14' 61 22 August 1997 58°03'—152°49' Depth ) (m) Substratum 257-265 silty sand, shingle 117-120 silty sand, shingle 150-164 silty sand, shell 180 silty sand Page 196 The Veliger, Vol. 43, No. 2 Figure 1 Pyrulofusus dexius (No 3/46838). A, B. Egg capsule; scale bar = 20 mm; C. A pattern on the outer surface of the egg capsule; scale bar = 1 mm. five wide, oblique, S-shaped axial folds appeared on the third whorl. The capsule and young specimens it con- tained were studied and described in vivo, after which they were preserved in formalin. Unfortunately, the shell of the young specimens dissolved in the formalin and it was impossible to take a photo of it. From the capsules of P. deformis and P. harpa (Gonor, 1964; Cowan, 1965), the capsule of P. dexius differs by its much greater diameter and flattened low-domed form. The young differ by the dextral shell and by the number of spiral ridges (Table 2). The egg capsule of P. melonus (Dall, 1891) remains so far unknown. Similar to P. dexius, it has a dextral shell, but these species differ fairly well by the shell sculpture, which is also clearly seen even in the capsule young. Literature Cited Cowan, I. M. 1965. The egg capsules and young of the gastropod Pyrulofusus harpa (Morch) (Neptuneidae). The Veliger 8:1—2. Gonor, J. J. 1964. Egg capsules and young of the gastropod Pyrulofusus deformis (Neptuneidae) at Barrow, Alaska. The Arctic 17:48—51. Hyman, L. H. 1967. The Invertebrates. Vol 6: Mollusca 1. Mc- Graw Hill: New York. 792 pp. KANTOoR, Yu.I. 1990. Gastropods of Subfamilia Volutopsiinae of the World Ocean. Nauka Press: Moscow. 180 pp. [In Rus- sian] THORSON, G. 1946. Reproduction and Larval Development of Danish Marine Bottom Invertebrates, with Special Reference to the Planktonic Larvae in the Sound (@resund). K@benhayn. 523 pp. Table 2 A comparison of the egg capsules and young of the three species of the North Pacific and subarctic Pyrulofusus. Characteristic P. dexius P. harpa P. deformis Capsule: Covering outer layers fine-patterned patterned without pattern inner layer smooth smooth ? Shape: low-domed hemispherical subspherical Maximum diameter 58 mm 39 mm 27 mm height 20 mm 21 mm 24 mm Young: Shell length 20.1 mm 19.5 mm 17.8 mm Spiral sculpture appears on third whorl appears on third whorl appears on second whorl 32 ridges 24+ ridges 40+ ridges Adult: Shell form dextral sinistral sinistral Maximum height 151.2 mm 122 mm 144.4 mm Information for Contributors Manuscripts Manuscripts must be typed, one side only, on A4 or equivalent (e.g., 842” X 11”) white paper, and double-spaced throughout, including references, figure legends, footnotes, and tables. All margins should be at least 25 mm wide. Text should be ragged right (ie., not full justified). Avoid hyphenating words at the right margin. Manuscripts, in- cluding figures, should be submitted in triplicate. The first mention in the text of the scientific name of a species should be accompanied by the taxonomic authority, in- cluding the year, if possible. 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Submitting manuscripts Send manuscripts, proofs, books for review, and corre- spondence on editorial matters to Dr. Barry Roth, Editor, 745 Cole Street, San Francisco, CA 94117, USA. CONTENTS — Continued Two new species of the family Hydrobiidae (Mollusca: Caenogastropoda) from Austria MARTIN HAASE, ERICH WEIGAND, AND HARALD HASEKE ............0.00.0000000- W9. Additional data on the phanerobranch dorid Zambja simplex Ortea & Moro, 1998 (Gastropoda: Nudibranchia: Polyceratidae) JUAN LUCAS CERVERA, JOSE CARLOS GARCIA-GOMEZ, AND RICCARDO CATTANEO-VIETTI .. 190 NOTES, INFORMATION & NEWS The egg capsule and young of the gastropod Pyrulofusus dexius (Dall) (Buccinidae: Volutopsiinae) VIEADIMIR: VEGUBBING Sisco, 5 eo ous ae iyo ss 5, StSdR aan oeeemene os ase ReL Senn CERN ee 195 VELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler, Founding Editor Volume 43 July 3, 2000 CONTENTS Revision of dorid Nudibranchia collected during the French Cape Horn Expedition in 1882-1883, with discussion of the genus Geitodoris Bergh, 1891 IMIT@EVAB IES CHIR I lepres aresetesy o aap waa peat ou ne MID EN On RUE ERE eS Sold. Sa 30h, 456 Sex change in the hat snail, Calyptraea morbida (Reeve) (Gastropoda: Calyptraeidae): an analysis of substratum, size, and reproductive characteristics MUN G= Awe CEENPANDIIGERVEAGS OON Garant oimaeiasiare aetate-ne tee tr tenis nines Sire tes oe Helicarionid snails of Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar IMENNEDEGC) PEMBERTON AND IMOTHY Av PEARCE ¢)...5. 2 e004 esc sates. Charopid snails of Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with description of a new genus and with conservation statuses of nine species KENNETH © EMBERTON AND DIMODHY A. PEARCE «6 c¢ sca cs bye ca ea oe ns Exploration of morphospace using Procrustes analysis in statoliths of cuttlefish and squid (Cephalopoda: Decabrachia)—evolutionary aspects of form disparity JEAN-LOUIS DOMMERGUES, PASCAL NEIGE, AND SIGURD V. BOLETZKY ...........-.- Shell size variation and aggregation behavior of Littoraria flava (Gastropoda: Littorinidae) on a southeastern Brazilian shore Pees PAWL 2 Bers a MOULINEO AND: CECINA PYALVES-COSTAY «se seen esse ane ec sais com ENMMIHSONTAR Jl n 19 2000 } LIBRARIES_— CONTENTS — Continued ISSN 0042-3211 197 210 218 248 265 The Veliger (ISSN 0042-3211) is published quarterly in January, April, July, and October by the California Malacozoological Society, Inc., % Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. Periodicals postage paid at Berkeley, CA and additional mailing offices. 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THE VELIGER © CMS, Inc., 2000 The Veliger 43(3):197—209 (July 3, 2000) Revision of Dorid Nudibranchia Collected during the French Cape Horn Expedition in 1882-1883, with Discussion of the Genus Geitodoris Bergh, 1891 MICHAEL SCHRODL Zoologische Staatssammlung Miinchen, Miinchhausenstr. 21, 81247 Miinchen, Germany Abstract. Historical material of cryptobranch Doridoidea collected from south of Tierra del Fuego during the ‘‘Mis- sion Scientifique du Cap Horn 1882-1883” and originally assigned to Doris hispida d’Orbigny, 1837, Doris plumulata Couthouy in Gould, 1852, and Doris luteola Couthouy in Gould, 1852, has been examined anatomically. One specimen is identified as the common Magellanic species Diaulula hispida (d’Orbigny, 1837). Two specimens belong to the poorly known Geitodoris patagonica Odhner, 1926. This latter species is redescribed for the first time using additional material recently collected. Re-examination of type material of G. patagonica and of Geitodoris falklandica Odhner, 1926, confirms the common possession of enlarged prostate glands and of large, saclike mantle glands. Both species are regarded to be conspecific, with Geitodoris patagonica Odhner, 1926, as the valid name. A lectotype of G. patagonica has been designated. This species is compared with congeners; Discodoris mavis Marcus & Marcus, 1967, and Discodoris tema Edmunds, 1968, are transferred to Geitodoris Bergh, 1891, due to the possession of spatulate marginal radular teeth which is considered to be an autapomorphy of the genus. This record from Nassau Bay and the synonymization with G. falklandica extends the known range of G. patagonica from Argentina to southernmost Chile and the Falkland Islands. The enigmatic species Doris plumulata and Doris luteola are discussed and regarded as nomina dubia. INTRODUCTION During the “Mission Scientifique du Cap Horn 1882-— 1883” several nudibranchs were collected from Nassau Bay, south of Tierra del Fuego, Chile, and tentatively identified by Rochebrune & Mabille (1891). All but Mi- crolophus poirieri Mabille & Rochebrune, 1891 (Den- dronotoidea) and Phidiana patagonica (d’ Orbigny, 1837) (Aeolidoidea) were doridoideans. The phanerobranch dorid Acanthodoris vatheleti Mabille & Rochebrune, 1891, was only briefly described externally by Rochebru- ne & Mabille (1891). Pruvot-Fol (1950) described addi- tional phanerobranch specimens collected during the Cape Horn Expedition which had not been mentioned by Rochebrune & Mabille (1891) as Thecacera darwini Pru- vot-Fol, 1950. This species was redescribed in detail by Marcus (1959), and, in living condition, by Schrédl (1996b). Most problematic are the cryptobranch dorids from the Cape Horn Expedition which Rochebrune & Mabille (1891) identified as Doris hispida d’ Orbigny, 1837, Doris plumulata Couthouy in Gould, 1852, and Doris luteola Couthouy in Gould, 1852 without giving any description. Diaulula hispida (d’Orbigny, 1837) is a better known species which is recognizable externally by a median dor- sal ridge (d’Orbigny, 1835—1846; Odhner, 1926; Marcus, 1959; Schrodl, 1996b). In contrast, both D. plumulata and D. luteola are highly dubious species. Due to Gould’s (1852, 1856) poor original descriptions, their correct identification appears extremely unlikely. This study aims to clarify the taxonomy of crypto- branch dorids collected during the Cape Horn Expedition on the basis of museum material. From dissections of historical specimens, detailed anatomical descriptions are presented for Diaulula hispida and Geitodoris patagoni- ca; for the latter species, type material and recently col- lected specimens were also examined. Geitodoris pata- gonica is revised taxonomically and compared with con- geners. SPECIES DESCRIPTIONS Family DISCODORIDIDAE Bergh, 1891 Genus Diaulula Bergh, 1880 Type species: Diaulula sandiegensis (Cooper, 1863) Diaulula hispida (ad Orbigny, 1837) (Figures 1—3) Doris hispida @ Orbigny, 1837:188, pl. 15, figs. 4-6; Roche- brune & Mabille 1891:10. Trippa hispida (d’ Orbigny, 1837): Bergh 1898:527—530, pl. 30, figs. 30-36, pl. 31, figs. 1-3; Odhner 1926:76—78, figs. 55-58, pl. 3, figs. 40-41 (‘‘d’Orbigny, 1836’); Carcelles 1950:70; Carcelles & Williamson 1951:315 (“‘d’Orbigny, 1847”’). Diaulula hispida (d’Orbigny, 1837): Marcus 1959:50—S3, Figure 1 SEM photograph (unsputtered) of radular teeth of Diaulula his- pida (“Doris sp.”’, MNHN). Scale bar = 200 ppm. figs. 109-114; Schrédl 1996a:52—53; 1996b:27, pl. 3, fig. 18; 1997a:39. Material examined: One specimen from the National Museum of Natural History (MNHN) of Paris labeled: Doris sp., Baie d’Orange, Baie de Nassau, Chili. Mission du Cap Horn (entrée 1883). External morphology: The single museum specimen is yellow, the foot darker than the notum. It is well extended and measures 23 mm in length, 12 mm in breadth, and 10 mm in height. The foot is broad, measuring 10 mm. However, this specimen is very poorly preserved. The mantle rim is seriously damaged so it might have been broader, and the anterior body is artificially swollen. Most parts of the notum are devoid of any recognizable tuber- cles, superficial tissue is almost completely destroyed. In a protected area lateral to the gills there are remains of small conical tubercles with diameters between 0.1 to 0.3 mm. There are about six multipinnate gills surrounded by an elevated sheath. The rhinophores are too damaged to give information on the number of perfoliations or the presence of a sheath. The foot is bilabiate anteriorly, but damaged near the mouth opening; the superior lip appears to be notched. The anterior mantle rim and the anterior part of the head is lacking, but there appears to be one digitiform oral tentacle left. Anatomy: Because the external features of this crypto- branch dorid do not allow identification, dissection was necessary. Owing to its very frail and amorphous consis- tency, my main attempt was to get information on cutic- The Veliger, Vol. 43, No. 3 hd qm a Se Q a 1mm d Figure 2 Reconstructed reproductive system of Diaulula hispida (“‘Doris sp.’, MNHN). Scale bar = 1 mm. Key: am, ampulla; bc, bursa copulatrix; fgm, female gland mass; hd, hermaphroditic duct; id, insemination duct; nd, nidamental duct; pb, penial bulb; pp, pe- nial papilla; pr, prostate; rs, receptaculum seminis; va, vagina; vd, vas deferens; vs, vas deferens sheath. ularized structures like lip cuticle, radula, and possible genital armature. The oral tube and pharynx are squeezed downward by an overlying swollen granular mass containing glandular particles, probably remains of the blood gland and sali- vary glands. Sand granules indicate the presence of the completely amorphous esophagus. Remains of the central nervous system could not be detected. Digestive system. The strong lip cuticle is brownish and smooth. The buccal mass is dark brown and hardened. It is dorsoventrally flattened, 6 mm in length, and 5 mm in breadth. The radula (Figure 1) measures 5.3 X 5.0 mm. It consists of 24 rows with up to 27 teeth per half row. The rhachis is small and lacks a central tooth. All lateral teeth are brownish, simple hamate without denticulation, and rather erect in shape with a blunt tip. The inner lat- erals are small, increasing in size toward the middle of the half rows (up to 0.3 mm). The outermost laterals are small. Reproductive system (Figure 2). Parts of the small con- ical penial papilla and the penial bulb were everted through the genital opening. The penis was dissolved in 10% KOH and did not possess cuticular armature. Inter- nally, the anterior genitalia are conglomerated and strong- ly hardened due to preservation. They are limited to the right side of the body. The ampulla is a curved tube at- tached to the large female gland mass. This organ appears to be composed of a dark, widely lobed mucus gland and a whitish, more granular albumen gland. There appears M. Schrédl, 2000 Page 199 Argentina Buenos Aires Juan Fernandez Islands = se wer tte eo ] Literature ©@ This study Figure 3 Geographic distribution of Diaulula hispida. Page 200 to be a large irregularly shaped prostate passing into an artificially squeezed, looped vas deferens section. The vas deferens consists of a narrow central duct which is sur- rounded by a thick muscular sheath. Distally the central duct passes into the small penial papilla; the muscular sheath fuses with the vestibular wall. There is a long, narrow vagina which widens distally into a vestibule sep- arate from the male opening. A bursa copulatrix and a receptaculum seminis were present but completely flat- tened. Their poor condition allowed a reconstruction of their vaginal arrangement, but not, with any certainty, their shape. Vagina and distal vas deferens were treated with KOH; both lack any cuticle. Discussion: The specimen examined within the present study was damaged and in very poor preserved condition. Internally, it fits with the description of Diaulula hispida (d’Orbigny, 1837), a species which was redescribed in detail by Marcus (1959) and recently, in living condition, by Schr6édl (1996b). In particular, radular features like the number of rows, teeth per half row, and erect, simple hamate shape, and smooth lip cuticle; and also genital characters like the special arrangement of seminal recep- tacles, separate vestibules, and vas deferens proximally forming a prostate and distally a narrow duct which is surrounded by a thick muscular sheath agree perfectly with Marcus’ redescription. The general body shape and coloration, a relatively broad, bilabiate, and notched foot, the dense, small notal tubercles, and the elevated gill sheath confirm the identification as D. hispida. The pres- ence of an undulating dorsal crest, the most distinctive external character of D. hispida, however, could be nei- ther confirmed nor denied; the museum specimen was in too poor condition, especially the central parts of the no- tum. The only known congener from Chilean and Argen- tinian waters, Diaulula vestita (Abraham, 1877) clearly differs in lacking a notal crest (see Odhner, 1926; Marcus, 1959); the latter species should be critically compared with Anisodoris punctuolata (d’Orbigny, 1837). Diaulula hispida was recently found in the Magellan Strait south of Punta Arenas (Schr6édl, 1996a, b) near the collecting locality of the museum specimen, and has a wide Magellanic distribution (Schrédl, 1996b, 1997a; this study, see Figure 3). The record from Tumbez, Peru, by Carcelles (1950) and Carcelles & Williamson (1951) is not based on original data and was regarded as an error (Schrodl, 1996b). Rochebrune & Mabille (1891) mentioned the crypto- branch species Diaulula hispida (as Doris hispida) as be- ing found during the Cape Horn Expedition. It is, how- ever, not clear if this identification really referred to this specimen, which is now too damaged to show the char- acteristic notal crest, or if this specimen formerly was assigned to Doris plumulata or Doris luteola, the other two cryptobranch species mentioned by Rochebrune & Mabille (1891). There is neither information within that The Veliger, Vol. 43, No. 3 publication nor on the museum’s labels indicating which specimen each of the names applies to. This makes no difference in the case of D. hispida since this species is clearly identified in the present study. It will always re- main problematic regarding the specimens assigned to Doris plumulata and Doris luteola. Both species origi- nally were described only externally and in a sketchy way on board ship by Couthouy and were subsequently estab- lished in the publications of Gould (1852, text; 1856, drawings). Millen et al. (1994) considered external de- scriptions of central Chilean species by Gould to be in- adequate for re-identification. This is also true for the southern species Doris plumulata and Doris luteola: D. plumulata comes closest to Anisodoris punctuolata (d’Orbigny, 1837) due to its fine notal tuberculation and eight delicate tripinnate gills. Having nine pairs of lan- ceolate, simply pinnate plumules Doris luteola more re- sembles Gargamella immaculata Bergh, 1894, which possesses eight to 12 bi- or tripinnate gills (Schrédl, 1996b, 1997b). However, there are some other Magellan- ic cryptobranch species with small notal tubercles and digitiform oral tentacles, i.e., two Geitodoris species de- scribed by Odhner (1926), which would also fit the su- perficial external descriptions of both of these species; there is no information on diagnostic internal organs, i.e., radula and genitalia. Since according to Johnson (1964; personal communication) no type material has been ever designated for Doris plumulata (misspelled **Doris plan- ulata”’ by Carcelles (1950) and “Doris plunulata’’ by Carcelles & Williamson, (1951) and Doris luteola (mis- spelled ‘‘Doris lucteola’’ by Carcelles (1950) and Car- celles & Williamson, (1951), both species cannot be re- identified and are considered to be nomina dubia. Family DIscoDORIDIDAE Bergh, 1891 Genus Geitodoris Bergh, 1891 Type species: Geitodoris complanata (Verrill, 1880) Geitodoris patagonica Odhner, 1926 (Figures 4—9) Geitodoris patagonica Odhner, 1926:80—83, figs. 59-63, pl.3, figs. 42-43; Carcelles 1950:70; Carcelles & Wil- liamson 1951:315; Schrédl 1996b:57, pl.3, fig. 17. Geitodoris falklandica Odhner, 1926:83-85, figs. 64—69, pl. 3, figs. 44-46. Material examined (see Table 1): Two specimens from the Muséum National d’Histoire Naturelle, Paris (MNHN) labeled: Doris sp., Baie d’Orange, Baie de Nas- sau, Chili. Mission du Cap Horn (entrée 1883). The larger dissected specimen has been re-examined; the smaller specimen has been dissected and is described anatomi- cally. Eight specimens of Geitodoris patagonica Odhner, M. Schroédl, 2000 Page 201 R loom Figure 4 Drawings of Geitodoris patagonica. A. Living recently collected specimen No. 2. B. Damaged smaller museum specimen “Doris sp.”, NMNH. Scale bars = 1 cm. 1926, from Bahia Camarones, Argentina, collected by S. Gigglinger and M. Schrodl, 9 January 1995, at 2 to 12 m depth, most on macroalgae, using SCUBA. A photo- graph of a living specimen was given by Schrédl (1996b). Two specimens have been examined anatomically and de- posited as voucher specimens in the Zoologische Staats- sammlung Mtinchen (ZSM) under the numbers 19971031 and 19971032. Geitodoris falklandica, holotype from the Swedish Mu- seum of Natural History, Stockholm (SMNH, type col- lection 2304), collected at Stanley Harbour, Falkland Is- lands. Geitodoris patagonica, lectotype, SMNH type collec- tion 2306, collected at Puerto Madryn, Argentina, 23 Jan- uary 1896. Geitodoris patagonica, four specimens, three of them partly dissected, SMNH type collection 1016, collected at Puerto Madryn, Argentina, 9 January 1895. External morphology (Figure 4): The larger specimen from the MNHN (‘‘Doris sp.’’) is whitish and measures about 60 mm in length, 30 mm in width, and 15 mm in height. Beside being seriously damaged and hardened due to preservation, the anterior part of the body had been partly dissected by a former worker. The notum is cov- ered with different-sized tubercles, the largest reaching a diameter of 1 mm. The tubercles are slightly elevated, rounded knobs. Spicules are absent, most probably due to preservation. The notum is squeezed and compact in the central parts. Laterally it has a more spongy consis- tency with several sac- or bottlelike, hollow structures within (see Figure 5). These reach about 0.5 mm in di- ameter and probably are large subepidermal glands. Some clearly open onto the notal surface, and it appears that these openings are not preservation artifacts. There are seven mainly bipinnate gills around the elevated anal pa- pilla. Gills and rhinophores are surrounded by consider- ably elevated sheaths covered with small tubercles. The foot is broad and anteriorly bilabiate. The upper lip is notched. Oral tentacles are long and digitiform. The smaller MNHN specimen (‘‘Doris sp.””) measures 37 mm in length, 19 mm in width and about 10 mm in height. It is moderately extended, but in rather bad ex- ternal condition (Figure 4B). Where undamaged, the no- tum is covered with different-sized tubercles. The largest ones reach a diameter of 1 mm and are surrounded by smaller tubercles with usually 0.1 to 0.3 mm diameter. All are knobby and most are only slightly elevated. Spic- ules are absent, but cavities within the tubercles and the notum indicate that spicules were present in the living specimen. Large, saclike subepidermal glands as de- scribed above are present throughout the notum (Figure 5C). Far posteriorly there is an unnaturally expanded, prominent anal papilla surrounded by seven or eight mainly bipinnate gills. The most posterior part of the gills and the notum is damaged and turned to the ventral side. The gills, as well as the rhinophores, are surrounded by elevated sheaths which bear different-sized tubercles re- sembling those of the notum. The perfoliate rhinophores possess about 17 lamellae. The foot is nearly as broad as the notum. Anteriorly it may be bilabiate, but this portion is damaged. No information can be given on the presence of a notch or on the shape of oral tentacles. As in the larger specimen, genital openings are within the anterior third of the body on the right side. The tip of the penial papilla and parts of an everted penial sheath are visible. Anatomy: The anterior portion of the larger specimen from the MNHN was dissected by a former worker. The radula was removed from the specimen but not retained in the museum lot. The anterior genitals are still mainly in situ, but strongly hardened due to preservation and Page 202 The Veliger, Vol. 43, No. 3 Figure 5 SEM photographs of critical point dried notum structures of Geitodoris patagonica. Note the saclike, large subepi- dermal glands (arrows). A. Holotype G. falklandica, overview of a notum section. Scale bar = 0.5 mm. B. Holotype G. falklandica, notum section in detail; see also spaces of dissolved spicules (arrow-heads). Scale bar = 0.2 mm. C. Doris sp. smaller specimen, MNHN. Scale bar-= 0.5 mm. D. Lectotype of G. patagonica. Scale bar = 0.5 mm. hardly suitable for a detailed examination. However, the anatomy of the larger specimen appears to generally agree with that of the smaller specimen which is de- scribed in the following section: Digestive System. The oral tube is wide and flattened, the pharynx is artificially squeezed backward. The yel- lowish to brownish lip cuticle is covered by small simple rodlets. The radula of the smaller specimen comprises 25 rows. The rachis lacks a rachidian tooth and is very nar- row with innermost lateral teeth of apparently alternating rows being close together. There are up to 24 lateral teeth lacking any denticulation, but their shape strongly varies and additionally depends on the angle of view. By light microscopy, the first and second laterals are small, having a stout hook. The following laterals increase in size be- coming more erect. The outer laterals decrease, the out- M. Schrédl, 2000 Page 203 Figure 6 SEM photographs of radular structures of Geitodoris patagonica. A. Smaller MNHN specimen, older lateral teeth (unsputtered). Scale bar = 0.2 mm. B. Recently collected specimen No. 1, lamellate marginal and hamate outer lateral teeth. Scale bar = O.1 mm. ermost becoming spoonlike in appearance. In older rows only rudiments of marginal teeth were detected. In youn- ger rows there are up to about 15 marginal teeth, which are delicate, very close standing lamellae. Some are Figure 7 Reproductive system of Geitodoris patagonica (“Doris sp.”’, smaller specimen, MNHN). Scale bar = 2 mm. Key: am, am- pulla; be, bursa copulatrix; fgm, female gland mass; nd, nida- mental duct; pp, penial papilla; pr, prostate; rs, receptaculum seminis; va, vagina; vg, vaginal gland; vd, vas deferens. slightly fringed, which may be an artifact. By SEM ex- amination, the shafts of most laterals in older rows prove to be laterally flattened. Inner laterals have an arrow- headlike tip (Figure 6A). In younger rows, lateral teeth are more slender and hook-shaped. Lateral and marginal teeth resembling those of younger rows of the museum specimen were present in recently collected G. patagon- ica specimens (Figure 6B). In the smaller MNHN specimen the esophagus is a round tube which is looped twice before passing through the circumesophageal nerve ring. The salivary glands are long thin tubes. The esophagus curves ventrally, pene- trating the digestive gland. The stomach is completely covered by the digestive gland; only a bulbous caecum reaches its dorsal surface posterior to the intestine. Ad- jacent to the surface of the digestive gland the intestine runs to its anterior edge and curves back to the anal pa- pilla. Genital System (see Figures 7, 8). The gonads cover the digestive gland. A thin hermaphroditic duct widens into a flattened, curved, white ampulla. Before entering the female gland mass, the prostate arises, apparently di- rectly from the ampulla. The prostate continuously wid- ens into a huge, massive U-shaped organ which is closely attached to an ample bursa copulatrix. The proximal por- tion of the prostate is yellowish and homogenous, but more distally it is yellow and appears granular. Distally the prostate narrows abruptly giving rise to the narrow, long, and highly convoluted muscular vas deferens. This duct passes into an unarmed conical penial papilla sur- Page 204 The Veliger, Vol. 43, No. 3 pr DC fgm 5 ) VQ vd oF O \\ = B Imm DD Figure 8 Reproductive system of recently collected Geitodoris patagonica No. 1. A. Jn situ. B. Schematical outline. Scale bars = 1 mm. Key: am, ampulla; bc, bursa copulatrix; fgm, female gland mass; id, insemination duct; nd, nidamental duct; pb, penial bulb; pp, penial papilla; pr, prostate; rs, receptaculum seminis; va, vagina; vg, vaginal gland; vd, vas deferens. rounded by a swollen penial sheath. The vagina opens into a common vestibule close to the male opening. Very near to its opening the vagina bears an unarmed tubular, flattened, and convoluted gland. The vagina is long and leads directly to the bursa copulatrix. This is an oval or- gan with a maximum dimension of 10 mm and is filled with a yellowish compact mass. The bursa is serially in- serted by the convoluted vaginal duct. Far distally, before entering the female gland mass, this duct gives rise to the short stalk of the receptaculum seminis. This somewhat oval to pear-shaped organ reaches 2 mm in dimension. The female gland is a compact, strongly hardened organ which obviously consists of a widely lobed mucus gland partly surrounded by other, more granular appearing por- tions. The nidamental duct is short, has a swollen bulb, and opens separately. Further Organ Systems. The blood gland consists of two lobes, a posterior one covering the posterior portions of the central nervous system, and an anterior one. The cerebropleural ganglia are completely fused. The eyes ap- pear to be sessile. Taxonomic discussion of Geitodoris patagonica: As pointed out above, it is not clear to which species Roche- brune & Mabille (1891) originally assigned the two mu- seum specimens examined. In the present study, both specimens are regarded to be conspecific, although only the smaller specimen could be examined in detail. The lip cuticle having rodlets, the radula with simple hamate lateral and lamellate (= spatulate) marginal teeth, and the genital system having an unarmed penial papilla and a vaginal accessory gland indicate placement in the genus Geitodoris as reviewed by Miller (1996). The smaller mu- seum specimen in nearly all external and anatomical fea- tures agrees with the description of Geitodoris patagonica Odhner, 1926. It differs from the original description of G. falklandica Odhner, 1926, the second species of the genus known from Magellanic waters, due to the presence of large subepidermal glands. Both of these Magellanic species, however, are described as completely lacking a prostate (Odhner, 1926). In contrast, a large massive pros- tate is undoubtedly present in the MNHN material ex- amined. To clarify this discrepancy, eight specimens considered to be G. patagonica and briefly described in living con- dition (Schrédl, 1996b) have been re-examined, two of them internally (Figure 4A). These specimens, collected from the Patagonian coast of Argentina, externally and anatomically agree with the MNHN specimens, except for some obvious preservation artifacts like body damage and distortions and the dissolution of calcareous spicules. The somewhat larger and more elevated tubercles in the MNHN specimens may be due to their larger size (see Table 1); the shape of lateral radular teeth is not as var- iable as seen in the smaller MNHN specimen. The genital systems of all specimens examined fit the sketchy draw- ings of G. falklandica (Odhner, 1926:figs. 68, 69) with only one difference: at the position drawn, the vas defer- ens passes into the prostate, which is closely attached to the female glands and hard to distinguish from the female gland mass (Figure 8A). The prostate is very large and extends around the bursa copulatrix (Figures 7, 8). A similar, large prostate has also been found on re- examination of the partly dissected holotype of G. falk- landica. The type material of G. patagonica is composed of five smaller specimens, the larger ones extensively dis- Page 205 M. 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There is no originally designated holotype. A dis- tinct prostate is doubtlessly present in a rather well-pre- served specimen (SMNH No. 2306) which is here des- ignated lectotype of Geitodoris patagonica. The prostate covering the bursa copulatrix was found to be stored in a small vial together with the partly dissected specimen. It is concluded that Odhner misinterpreted the prostate as part of the female glands in both Geitodoris patagonica and G. falklandica. This might be due to the fact that the genus Geitodoris was clearly defined by radular criteria but believed to lack a distinct prostate at the time of Odhner’s study (see generic discussion: Odhner, 1926:78— 80). Owing to the presence of numerous subepidermal no- tum glands, the specimens examined during the present study agree with the original description and with type material of G. patagonica (see Figures 5C, D). In con- trast, G. falklandica was originally described as lacking such glands (Odhner, 1926). The holotype of G. falklan- dica indeed has a comparatively thin, somewhat squeezed notum. In lateral areas, however, there clearly are large saclike glands within the notum (see Figures 5A, B). The quantity of glands present in certain areas of the notum varies intra-individually and between recently collected specimens from the same population and is strongly in- fluenced by preservation (personal observation). Thus there remains no reason to doubt the conspecifity of G. patagonica and G. falklandica. Both species were estab- lished within the same study; priority is given to G. pa- tagonica Odhner, 1926, due to its name better reflecting the geographical distribution of the species and to its orig- inal description giving more details on the characteristic notum consistency. A single specimen from Puerto Que- quén, Argentina was assigned to G. patagonica (as “‘Gei- todoris patagonicus’’) by Carcelles (1944) with some res- ervations. Since no description was given, this record needs to be confirmed. Geographical distribution: Geitodoris patagonica was known previously from Puerto Madryn (about 42°30’S), Argentina (Odhner, 1926) south to Bahia Camarones (44°53'S, 65°39'W) (Schrédl, 1996b). The identification of material collected during the French Cape Horn Ex- pedition in 1882-1883 as G. patagonica extends this range south to Orange Bay, Hoste Island, Chile (Figure 9). The synonymy with G. falklandica includes the Falk- land Islands. In contrast to Diaulula hispida and several other Magellanic nudibranchs which occur on both the Atlantic and the Pacific coasts of Patagonia (Schrédl, 1996a, b, 1997a, b, c; this study, Figure 3), G. patagonica at the moment appears to be limited to Atlantic waters south to Tierra del Fuego and adjacent islands having a Falkland Current-related distribution (Figure 9). Howev- er, this picture may change with a better faunal knowl- edge of the southern Chilean fjord region and improved taxonomy. The Veliger, Vol. 43, No. 3 Generic comparison: In recent reviews (Ortea & Balles- teros, 1981; Ortea, 1990; Miller, 1996) the genus Geito- doris was mainly characterized by the presence of spat- ulate (lamellate, platelike) marginal radular teeth, in ad- dition to, but clearly differing from, more or less hook- shaped lateral teeth. The presence or absence of a distinct prostate has been used as an important character to dis- tinguish species within the genus Geitodoris as well as to define subgenera. Four species with smooth lateral teeth and without a differentiated prostate (G. complanata (Verrill, 1880); G. immunda Bergh, 1894; G. mollina Bergh, 1904; and G. lutea Baba, 1937) were placed into the subgenus Geito- doris Bergh, 1891 (type species: G. (G.) complanata) by Ortea & Ballesteros (1981). However, a curved ‘‘pros- tate’’ was explicitly mentioned in the original description of G. immunda by Bergh (1894), and Baba (1937) did not describe the genitalia of G. lutea at all. Miller (1996) added G. reticulata Eliot, 1906, to the subgenus Geito- doris sensu stricto. Curiously, Cervera et al. (1985) pre- viously considered G. reticulata to be a junior synonym of G. planata (Alder & Hancock, 1846), a species de- scribed as having a distinct prostate in the same study. In addition, Geitodoris patagonica and (its synonym) G. falklandica were assigned to the subgenus Geitodoris s. s. by Miller (1996). However, since G. patagonica is shown herein to possess a distinct prostate, this species must be re-compared with congeners having a prostate. Species assigned to the subgenus Carryodoris Vayssi- ere, 1919 (type species: G. (C.) joubini) by Ortea & Bal- lesteros (1981), Geitodoris joubini (Vayssiére, 1919), G. oshimai (Baba, 1936), and G. portmanni (Schmekel, 1970) have serrate lateral teeth and clearly differ from G. patagonica, which has simple hooked laterals. The pres- ence of a prostate in Carryodoris assumed by Ortea & Ballesteros (1981), however, was only confirmed for G. portmanni (see Schmekel, 1970; Schmekel & Portmann, 1982; Perrone, 1984). Geitodoris patagonica agrees with species of the sub- genus Verrillia Ortea & Ballesteros, 1981 (type species: G. (V.) bonosi), which according to Ortea & Ballesteros (1981) and Miller (1996), are characterized by a distinct prostate and smooth laterals. In Geitodoris bacalladoi Or- tea, 1990, and G. sticta Miller, 1996, a vestibular gland is absent (Miller, 1996), whereas G. patagonica clearly possesses a tubular vaginal gland (Odhner, 1926; this study). Geitodoris bonosi Ortea & Ballesteros, 1981, was said to have a penial gland, but figure 3 of the same paper (Ortea & Ballesteros, 1981) shows a gland opening as close to the vagina as to the male duct. In contrast to G. patagonica, which has a receptaculum seminis on a short stalk, G. bonosi is described as having a serially arranged receptacle (Perrone, 1992). Geitodoris perfossa Ortea, 1990, G. capensis Bergh, 1907, G. planata (Alder & Han- cock, 1846), and G. pusae (Marcus, 1955) are all distin- guishable from G. patagonica due to their yellow, orange, M. Schrédl, 2000 Page 207 wreorrn Argentina Buenos Aires Juan Fernandez Islands Island Literature @ This study Figure 9 Geographical distribution of Geitodoris patagonica. Page 208 reddish, or brownish coloration with white and brownish patches on the notum (see Ortea, 1990; Miller, 1996). Geitodoris patagonica has a whitish notum with irregular black spots (Schrédl, 1996b). Geitodoris pusae is unique in having spicules (Ortea et al., 1988; Ortea, 1990) or cuticular spines (Marcus & Marcus, 1967) within the dis- tal portion of the vestibular gland. Geitodoris immunda, never placed in Verrillia, although possessing a prostate (Bergh, 1894), has serrate marginal teeth, whereas the marginals are smooth or irregularly worn in G. patagon- ica. Miller (1996) discussed four additional species as pos- sibly belonging to the genus Geitodoris, but marginal rad- ular teeth remain to be confirmed as lamellate in Disco- doris palma Allan, 1933, Discodoris crawfordi Burn, 1969, and Discodoris millegrana (Alder & Hancock, 1854). Only one species, Geitodoris heathi (MacFarland, 1905) resembles G. patagonica due to the possession of lamellate marginal teeth, and of prostate, vestibular, and saclike mantle glands (Marcus, 1961; Millen, personal communication). All the above species differ from G. pa- tagonica in having orange or brownish pigmentation. Discodoris mavis Marcus & Marcus, 1967, a species very similar to G. heathi, is transferred to the genus Gei- todoris herein due to its spatulate marginal teeth. It re- sembles G. patagonica in having a swollen prostate, a vaginal gland, a stalked receptaculum seminis and saclike mantle glands, but differs in its orange coloration (Marcus & Marcus, 1967). Discodoris tema Edmunds, 1968, from Ghana is also transferred to the genus Geitodoris. This is due to the possession of close-standing, spatulate margin- al teeth (see Edmunds, 1968:fig. 7) clearly differing in shape from the simple hamate laterals. This orange spe- cies externally differs from G. patagonica, but closely resembles G. perfossa Ortea, 1990. In conclusion, G. pa- tagonica is a valid species and does not appear to have any synonyms beside G. falklandica. Within nudibranchs close-standing, spatulate marginal radular teeth are restricted to members of the genus Gei- todoris. Presuming this unique character to be derived from uniform, hamate teeth (Gosliner, 1994) only once, the genus Geitodoris would be a monophyletic group. In contrast, the subgenera established by Ortea & Ballester- os (1981) were defined on the base of character combi- nations. Therefore, they do not necessarily reflect natural groups, and future cladistic analysis would be desirable. Acknowledgments. My thanks go to Phillippe Bouchet (MNHN) for kindly allowing me to dissect museum specimens, as well as to Anders Warén (SMNH) for providing type material for re- examination. David Reid (BMNH), Bill Pettitt (Manchester Mu- seum), and Richard I. Johnson (USNM) are thanked for their help in tracing historical material. Sandra Millen (Vancouver) kindly gave me unpublished data on G. heathi. Heike Wigele (Bochum) and Sandra Millen are acknowledged for helpful com- ments On an earlier version of the manuscript. Sebastian Gig- glinger helped me during field work, which was financed by grants of the Deutsche Akademische Austauschdienst (DAAD). The Veliger, Vol. 43, No. 3 LITERATURE CITED BaBA, K. 1937. Opisthobranchia of Japan. Journal of the De- partment of Agriculture, Kyushu Imperial University 6:1— 19. BeRGH, L. S. R. 1894. Die Opisthobranchien. Bulletin of the Museum of Comparative Zoology Harvard 25:125—235, pls. 1-12. BerRGH, L. S. R. 1898. Die Opisthobranchier der Sammlung Plate. Zoologische Jahrbiicher Supplement 4:48 1—582. CARCELLES, A. R. 1944. Catalogo de los moluscos marinos de Puerto Quequén. Revista del Museo de La Plata N. S. Sec- cidn Zoologia 3:233-309, pls. 1-15. CARCELLES, A. R. 1950. Catalogo de los moluscos marinos de la Patagonia. Anales Museo de Nahuel Huapi 2:41—99, pls. 1-6. CARCELLES, A. R. & S. I. WILLIAMSON. 1951. Catalogo de los moluscos marinos de la Provincia Magallanica. Revista del Instituto Nacional de Investigacion de Ciencias Naturales, (Ci. Zool.) Buenos Aires 2:225—383. CERVERA, J. L., J. C. GARciIA & F J. Garcia. 1985. Redescription of Geitodoris planata (Alder & Hancock, 1846) (Gastro- poda: Nudibranchia). Journal of Molluscan Studies 51:198— 204. D’ORBIGNY, A. 1835—46. Voyage dans 1’ Amérique Méridionale exécuté pendant les années 1826-1833. Vol. 5. Mollusques. Libraire de la Société geologique de France, Paris. 758 pp. plus Atlas. EDMUNDS, M. 1968. Opisthobranchiate Mollusca from Ghana. Proceedings of the Malacological Society of London 38:83— 100. GOSLINER, T. M. 1994. Gastropoda: Opisthobranchia. Pp. 253— 355 in FE Harrison & A. J. Kohn (eds.), Microscopic Anat- omy of Invertebrates. Vol. 5: Mollusca I. Wiley-Liss: New York. GouLpb, A. A. 1852, 1856. United States Exploring Expedition During the Years 1838-1842. Mollusca & Shells 12:I-XV, 1-510, 1852 with an Atlas of plates, 1856. JOHNSON, R. I. 1964. The recent Mollusca of Augustus Addison Gould. Bulletin of the United States National Museum 239: 1-182, pls. 1-45. Marcus, E. 1959. Lamellariacea und Opisthobranchia. Reports of the Lund University Chile Expedition 1948-49, No. 36. Lunds Universitets Arsskrift N.F 55:1-133. Marcus, E. 1961. Opisthobranch mollusks from California. The Veliger 3(Suppl.):1—85, pls. 1-10. Marcus, Ev. & ER. Marcus. 1967. American Opisthobranch Mollusks. Institute of Marine Sciences: Miami. vii + 256 Pp- MILLEN, S. V., M. SCHRODL, N. VARGAS & A. INDACOCHEA. 1994. A new species of Okenia (Nudibranchia: Doridacea) from the Peruvian Faunal Province. The Veliger 37:312-318. MILLER, M. C. 1996. A new species of the dorid nudibranch genus Geitodoris Bergh, 1892 (Gastropoda, Opisthobran- chia) from New Zealand. Journal of Molluscan Studies 62: 433-442. Opuner, N. H. 1926. Die Opisthobranchien. Further Zoological Results of the Swedish Antarctic Expedition 1901—1903 2: 1—100. OrtTEA, J. 1990. El género Geitodoris Bergh, 1891 (Mollusca: Nudibranchia) en las Islas Canarias. Revista de la Academia Canaria de Ciencias 2:99—120. OrTEA, J. & M. BALLESTEROS. 1981. A new Doridacea from the Iberian and Balearic littoral: Geitodoris bonosi n. sp. Journal of Molluscan Studies 47:337-342. OrTEA, J., A. A. LUQUE & J. TEMPLADO. 1988. Elysia picta Ver- M. Schrédl, 2000 rill, 1901, and Geitodoris pusae (Marcus, 1955), two am- phiatlantic opisthobranch gastropods. Journal of Molluscan Studies 54:243-247. PERRONE, A. S. 1984. Contributo alla conoscenza di Geitodoris (Carryodoris) portmanni (Schmekel, 1970) (Opisthobran- chia: Nudibranchia). Bolletino Malacologico 20:139—150. PERRONE, A. S. 1992. Una specie di nudibranchi nuova per le coste italiane: ridescrizione di Geitodoris (Verrilia) bonosi Ortea & Ballesteros, 1981 (Opisthobranchia: Nudibranchia). Bolletino Malacologico 28:27—34. PruvoT-FoL, A. 1950. Le genre Thecacera Fleming 1828 et une espece nouvelle: Thecacera darwini. Journal de Conchylio- logie Paris 90:48-—52. ROCHEBRUNE, A. T. & J. MABILLE. 1891. Mollusques. Mission Scientifique du Cap Horn 6, Moll., Paris. 192 pp., 8 pls. SCHMEKEL, L. 1970. Eine neue Art der verschollenen Gattung Carryodoris Vayssiére, 1919 aus dem Golf von Neapel, Car- ryodoris portmanni n. sp. (Gastr. Nudibranchia). Pubblic- cacioni della Stazione Zoologica di Napoli 38:370—377. Page 209 SCHMEKEL, L. & A. PORTMANN. 1982. Opisthobranchia des Mit- telmeeres. Nudibranchia und Saccoglossa. Fauna e Flora del Golfo di Napoli, 40. Springer-Verlag: Berlin. 410 pp. SCHRODL, M. 1996a. Opisthobranchs (Gastropoda) collected by the research vessel ‘Victor Hensen.” Berichte zur Polarfor- schung 160:52—54. SCHRODL, M. 1996b. Nudibranchia y Sacoglossa de Chile: mor- fologia exterior y distribuci6n. Gayana Zoologia 60:17—62. SCHRODL, M. 1997a. Range extensions of Magellanic Nudi- branchs (Opisthobranchia) into the Peruvian Faunal Prov- ince. The Veliger 40:38—42. SCHRODL, M. 1997b. On the Magellanic nudibranch Gargamella immaculata Bergh, 1894, and its synonymy to G. latior Odhner, 1926. Spixiana 20:81—92. SCHRODL, M. 1997c. On the morphology of the Magellanic nu- dibranch Anisodoris fontaini (d’Orbigny, 1837), and its syn- onymy with A. tessellata Bergh, 1898. The Veliger 40:228— 233. THE VELIGER © CMS, Inc., 2000 The Veliger 43(3):210—217 (July 3, 2000) Sex Change in the Hat Snail, Calyptraea morbida (Reeve) (Gastropoda: Calyptraeidae): An Analysis of Substratum, Size, and Reproductive Characteristics MING-HUI CHEN anpD KERYEA SOONG! Institute of Marine Biology, National Sun Yat-sen University, Kaohsiung, Taiwan 804, Republic of China (e-mail: keryea@mail.nsysu.edu.tw) Abstract. The substratum, body size, and reproductive characteristics of the hat snail, Calyptraea morbida, from the west coast of Taiwan were studied. Among 416 snails attached to a substrate, 77.2% were found on the shells of Turritella terebra, which were usually occupied by the hermit crab, Diogene spinifrons. The shell margin of the female C. morbida was more irregular than that of the male. Moreover, C. morbida females on the shells of 7. terebra were significantly heavier than those on smaller host. shells (mostly Turricula javana and Nassarius clathratus). No such difference was found in males. Female snails were significantly heavier than males. Individuals smaller than 18 mg were all male, whereas those larger than 202 mg were all female. Females had a shorter penis (< 2 mm) than that of the males (> 2 mm). A significant negative correlation between penis length and snail size was found in females but not in males. Brooding individuals were present almost year-round, with the highest occurrence in May. The recruitment of males (< 25 mg) occurred year- round, with a peak in July. Females smaller than 75 mg, on the other hand, were recruited between January and September, with the highest relative frequencies occurring in April. The size difference between sexes and reduction of penis length in females suggests that a life history involving sex change occurs in C. morbida. INTRODUCTION Sex change or sequential hermaphroditism is a phenom- enon in which an organism functions first as one sex, then as another in the later stage. It is divided into three cat- egories: protandry (males change to females), protogyny (females change to males), and alternating sexuality (re- peated change of sex) (Warner, 1988; Wright, 1988; Hel- ler, 1993). Protandry is widely scattered among inverte- brates, while protogyny is very common among coral reef fishes (Policansky, 1982). Alternating sexuality might be rare in animals and it can be found in few species of bivalves and polychaetes (Coe, 1932; Policansky, 1982; Berglund, 1986; Heller, 1993; Premoli & Sella, 1995). Because sex change has evolved independently under many disparate circumstances, life history theory has sought to determine the conditions under which sex change is favored over either simultaneous hermaphro- ditism or gonochorism. A common explanation of sex change is the size (or age) advantage hypothesis, which predicts that sex change is favored when the reproductive success of the two sexes is different with respect to size (or age). For example, protandry may be favored when female egg production increases with size, while the abil- ity of males to fertilize eggs is independent of size (Ghi- ' Corresponding author selin, 1969; Warner, 1975; Charnov, 1982; Warner, 1988; Charnov & Bull, 1989; Collin, 1995). Protandry has been documented in several groups of marine mollusks, including the gastropod superfamilies Calyptracea and Patellacea, and the bivalve superfamily Galeommatacea (Hoagland, 1978; Policansky, 1982; Heller, 1993). Among these, the gastropod family Calyp- traeidae is the best known (Hoagland, 1978; Wright, 1988). All species in the calyptraeid genus Crepidula La- marck, 1799, which have been investigated to date are protandric (Coe, 1938; Hoagland, 1978; Heller, 1993; Warner et al., 1996). They attach themselves to hard sub- strata and are sedentary. Some species form large stacks consisting of large females, intersexed individuals, and small males [e.g., Crepidula fornicata (Linnaeus, 1758), and Cr. onyx Sowerby, 1824], while others form male- female pairs (e.g., Cr. convexa Say, 1822, Cr. plana Say, 1822). The timing of sex change might be influenced by the gender of conspecific neighbors (e.g., Cr. fornicata and Cr. onyx) (Coe, 1938; Hoagland, 1978; Warner et al., 1996). In contrast to Crepidula, sex change has been studied only in one species in the genus Calyptraea. The sex change of Calyptraea chinensis (Linnaeus, 1758) differs from that of Crepidula in several aspects. In C. chinensis sex change always occurs at a certain age in its life history (Pellegrini, 1949; Wyatt, 1961) and not M.-H. Chen & K. Soong, 2000 Page 211 Table 1 Calyptraea morbida. Sizes of males and females in each collection. Numbers in parentheses indicate sample sizes. Body weight! (mg) Year Month Male 1991 April 40 (261) May 15 (15) July 27 (178) August 42 (133) November 26 (5) December 48.5 (32) 1992 January 39 (176) March 61.5 (144) May 32 (656) September 63 (45) November 74 (27) Combined SHES (1672) Female 140 (242) 170 (7) 160 (53) 146 (62) 110.5 (8) 166 (18) 129.5 (32) 177 (52) 134 (327) WA (17) 204 (15) 142 (833) Male only Overlapping range Female only Sex ratio? range (m, f) range (male/all) 4-34 35-200 208-735 O'S2"8 (114) (147, 189) (53) 8-47 48—48 52-223 0.68" (14) (QE IY) (6) 1-44 45-140 150-518 OW (128) (50, 19) (34) 6-58 59-136 138-596 0.68** (91) (42, 23) (39) 8-55 no 60-295 0.38" (5) (8) 10-79 80-113 115—490 0.64* (29) (3, 3) (15) 7—46 47-148 160-347 0.85** (111) (65, 20) (12) 8-57 58-162 165-342 OW (60) (84, 23) (29) 1-17 18-194 195-388 0.67** (126) (530, 262) (65) 9-4] 43-146 148-324 0.73** (15) (30, 5) (12) 4-95 102—202 204—468 0.64" (18) Q, 7) (8) 1-17 18-202 203-738 0.66** (248) (2068) (189) "Median of body weight. All comparisons of body weight between males and females were significant (P < 0.01, Mann-Whitney U- test). > x?-test for sex ratios deviating from 1:1. ™: P > 0.05; *: P < 0.05; **: P < 0.01. If a sequential Bonferroni technique for correction of multiple tests (Lessios, 1992) is applied, seven tests, except the one from December 1991, remain significantly biased from 1:1. at different times in different animals, as in Crepidula. Calyptraea chinensis breeds between December and May in Naples, Italy, with male recruitment occurring there- after; and sex change occurs from January to March (Pel- legrini, 1949; Bacci, 1951). Males and females are asso- Gastropoda ciated only during the breeding season when the smaller males are carried by the females. There appears to be no self-fertilization (Wyatt, 1960). The hat snail, Calyptraea morbida (Reeve, 1859), has a limpetlike shell with a septum. It is distributed at depths Table 2 Occurrences of Calyptraea morbida on different substrata from west coast of Chadin, Taiwan. Substratum Babylonia formosae (Sowerby, 1866) Ficus ficus (Linnaeus, 1758) Nassarius clathratus naticids Turricula javana Turritella terebra Bivalves Total (%) * The hermit crab is Diogene spinifrons. Host shell Occupied by a Empty hermit crab* Total (%) (0) 14 15 (3.6) 0) 3 3 (0.7) 5 18 24 (5.8) Y 12 19 (4.6) 8 20 29 (7.0) 78 243 321 (77.2) 5 0) 5 (1.2) 103 (24.8) 310 (74.5) 416 (100.0) (b) Minimum Height Maximum Width Figure | Calyptraea morbida. Shell morphology. (a) Left side view of a female shell, (b) Left side view of a male shell, (c) Apical view of a female shell, (d) Apical view of a male shell. of 10-100 meters along the west coast of Taiwan and along the south-east coast of mainland China. It is found attached to mollusk shells housing hermit crabs and emp- ty mollusk shells on the bottom of soft sediment. No pub- lished information on the life history of C. morbida is available. Therefore, it was previously unknown if these snails change sex. In order to elucidate the possible sex change of C. morbida, the present study examined the substratum, body size, and reproductive characteristics of the hat snail as well as its life history. The Veliger, Vol. 43, No. 3 MATERIALS AnD METHODS Specimen Collection and Substratum Types Specimens of Calyptraea morbida were collected by bottom dredging at 10—60 meters depth using a mesh size of about 1.1 cm along the west coast of Taiwan near Chadin (22°40’N, 120°09’E). A total of 11 collections were made between April 1991 and November 1992 (see Table 1). The specimens were kept in crushed ice during transportation; this practice may contribute to the high frequency of snails detached from their substate. If the snails were still attached to the substrate upon examina- tion, both the snails and the substrate were kept in the same plastic bags. These specimens were fixed in 10% formalin and later preserved in 70% ethanol. The type of substrate (i.e., live mollusks, hermit crabs, and empty shells) was recorded for each hat snail. Since linear mea- surements of shells were not good indicators of size, the body weight without shell was used instead. After blot- ting with a piece of tissue paper, the wet body weight of each individual was measured on an electronic balance with a sensitivity of 1 mg. Some specimens in this study were deposited in the National Museum of Natural Sci- ences (serial# NMNS 002894) in Taichung, Taiwan. Shell Morphology The maximum and minimum height and width was measured for each shell (Figure 1) using a pair of vernier calipers for 90 specimens collected in April 1991. The ratio of minimum shell height to maximum shell height (RH) and the ratio of minimum shell width to maximum shell width (RW) were used to indicate the variability of the shell morphology which is mostly caused by substrate curvature and irregularities. Size Structure and Reproductive Characteristics All the specimens were sexed after dislodging the soft body from the shells. Females were distinguished by the presence of the white-colored capsule gland located on the right edge of the mantle, at the anterior of the colu- mellar muscle. The presence of egg capsules containing Table 3 Calyptraea morbida. Size comparison of snails on Turritella terebra and on other host shells. Body weight (mg) Sex Substratum Median Male Turritella terebra 45.5 Others* 39 Female Turritella terebra 179 Others 139 Range n Mann-Whitney U-test 1-180 178 P > 0.05 10—202 59 18-596 143 P< 0.01 43-439 36 “Including Babylonia formosae, Ficus ficus, Nassarius clathrata, naticids, Turricula javana, and bivalves. M.-H. Chen & K. Soong, 2000 Page 213 LOO 2S) July n=231 Frequency December n=50 LOZ 100 January q5 n=208 50 25 0 100 75 March aS 50 n=196 50 25 25 0 0 250 May n=983 200 150 100 50 0 November n=42 50 August n=195 15 September 25 10 n=62 0 5 0 100200300400500600700800 9 100 200 300 400 500 Body weight (mg) Figure 2 The size distributions of Calyptraea morbida in different months. The collections in May and November, 1991, with small sample sizes (n < 30), are not shown. embryos could only be recorded for those females still attached to the substratum because in detaching females, the egg capsules were left attached to the substratum. To- tal numbers of embryos in all capsules were counted in 12 individuals to assess the relationship between snail size and fecundity. The lengths of penes were measured under a stereomicroscope for all specimens collected in April 1991. A total of 31 hat snails (17 males and 14 females) were sectioned at 5 wm for histological gonad development study. Sections were stained with Harris hematoxylin and eosin. RESULTS Types of Substrata Of 2505 specimens examined, 2089 individuals (83.4%) were found detached from their substrate, and 416 individuals (16.6%) were still attached to a substrate (Table 2). Among those attached, 77.2% occurred on the %* Small males Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month Figure 3 Calyptraea morbida. The annual change of the relative frequen- cies of small males (< 25 mg). Numbers above bars indicate sample sizes. body whorl of shells Turritella terebra (Linnaeus, 1758). These host shells were mostly (74.5%) occupied by the hermit crab, Diogene spinifrons. Shell Morphology and Size Structure The shell of Calyptraea morbida is drawn as Figure 1. The shell margin of C. morbida was highly variable depending on the surface contour of the substratum. The RH and RW were used as indices to indicate shell var- iation. Seventeen out of 58 female shells and three out of 32 male shells had an RH value smaller than 0.76. Also, 15 out of 58 female shells and one out of 32 male shells had an RW value smaller than 0.80. In both of the above cases, the frequency of snails with extreme values of indices was dependent on the sex of the snails (Chi-square tests, P = 0.03 for RH, P < 0.01 for RW). Within the size range of 51-160 mg, in which individ- uals of both genders occur, females were found to have a significantly smaller RH (mean = 0.80, median = 0.79) than males (mean = 0.83, median = 0.83) (n = 58, P < 0.05 Mann-Whitney U-test). No such difference was found in RW. There is no significant correlation between RH or RW, and body weight in either males (7? = 0.0004 for RH, 7? = 0.0729 for RW, P > 0.05) or females (r* = 0.0036 for RH, r? = 0.0004 for RW, P > 0.05). Thus the variation in shell morphology cannot be attributed to growth. Since the shell margin was irregular, the relative size of the hat snail is expressed in term of the wet body weight. Females on the shells of Turritella terebra were significantly heavier than those on other substrata [mainly Turricula javana (Linnaeus, 1758) and Nassarius clath- ratus (Born, 1778)]. However, no such difference was found in males (Table 3). There was no size difference The Veliger, Vol. 43, No. 3 % Small females Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month Figure 4 Calyptraea morbida. The annual change of the relative frequen- cies of small females (< 75 mg). Numbers above bars indicate sample sizes. between the snails on empty shells and shells occupied by hermit crabs. In this study a total of 2505 individuals ranged from 1 to 735 mg in wet weight. The size of females was signif- icantly larger than that of the males in each of 11 samples (P < 0.01, Mann-Whitney U-test, Table 1). The size of males and females often overlapped, although in all col- lections the smallest animals were all males and the largest snails were all females (Table 1, Figure 2). After pooling all collections, the male-only range was 1-17 mg and the female-only range was 203-735 mg. The size range in- cluding both sexes was 18-202 mg, representing 25.2% of the total size range; and 82.6% of all individuals feel into this range. Males smaller than 25 mg, presumably recently recruited, were found in each collection throughout the study. The highest frequency of these recruiting males rel- ative to all snails was found in July (Figure 3). Females smaller than 75 mg, presumed to have changed from males recently due to their sizes, were found between January Table 4 Calyptraea morbida. The distribution of male and female snails on individual shells. No. of observa- No. of Group Type tions snails One-snail Male 148 n = 277 Female 129 Two-snail 2 males 17 n= 118 male-female 39 2 females 3 Three-snail 2 males, 1 female 5) n= 21 3 males 2 M.-H. Chen & K. Soong, 2000 9 8 1 z Mare Y=0.004xX+2.91 Ye 6 r7=0.01,n=261 © 59°° p=0.11 5 ° Reha Es 2 a4 D a 0 ~ 0 100 200 300 9 n ‘4d 8 ¢ S 7 Female 6 Y=-0.002X+1.75 5 ° r2=0.09,n=242 je p<0.01 3 2 1 0 0 100 200 300 400 500 600 700 800 Body weight (mg) Figure 5 The relationship between Calyptraea morbida body weight and the length of penes for all snails collected in April 1991. and September, with their highest frequency relative to all females appearing in April (Figure 4). Reproductive Characteristics Sex ratio: A total of 1672 males and 833 females were collected. The sex ratios of most (7/11) samples were significantly biased toward males, and none were fe- male-biased (Table 1). Among those hat snails still at- tached to the substrata, most were solitary (66.6%) and the remainder were in groups of two or occasionally three (Table 4). Histological examination and penis length: Among 17 males (range: 4-130 mg) examined histologically, the smallest male observed with sperm was 23 mg in wet body weight. Various stages of ovarian development were found in 14 sectioned females (range: 67—420 mg). None was seen with both sperm and oocytes together. Most females (86.0%, n = 242) had their penes shorter than 2 mm, whereas the length of penis in most males (86.6%, n = 261) was longer than 2 mm. A significantly Page 215 % Brooding Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month Figure 6 Calyptraea morbida. The annual change of the relative frequen- cies of brooding females. Numbers above bars indicate the total numbers of females that could be examined for the presence of egg capsules. *Brooding individuals were found, but the frequency was not available in the month. No collection available in February, June, and October. negative correlation between the length of penes and the size of snails was found in females but not in males (Fig- ure 5). Fecundity: The egg mass of an individual consists of a cluster of egg capsules each of which is connected by a filament to a sticky pad attached to the substratum. All embryos in a single brood were at the same stage of de- velopment. Females had brood at a variety of stages of development in every collection throughout the study pe- riod. The percentage of females with egg capsules was the highest in May (81%, Figure 6). Each female had 16— 43 egg capsules per brood (n = 12, female size range 123-327 mg) and each capsule had 6—71 embryos. Re- lease of planktonic larvae from the egg capsules was ob- served in the laboratory. Brooding females (median: 178 mg, range: 72-596 mg, n = 122) were significantly larger than non-brooding females (median: 150 mg, range: 18— 468 mg, n = 57) (P < 0.01, Mann-Whitney U-test). The smallest brooding female was 72 mg. Large females were more likely to have egg capsules than small ones (Figure 7). Moreover, the fecundity increases with size (r? = 0.40, P < 0.05, n = 12) (Figure 8). DISCUSSION Type of Substrate Calyptraeids often utilize shells occupied by hermit crabs as their substrata (Table 2) (Hendler & Franz, 1971; Hoagland, 1977a, b; Karlson & Cariolou, 1982; Shenk & Karlson, 1986; Vermeij, 1989). This association might in- crease the dispersal of males in calyptraeid species (Hen- Brooding % <50 50-100 100-150 150-200 200-250 >250 Body weight (mg) Figure 7 Calyptraea morbida. The percentage of females that are brooding in different size groups. Numbers indicate sample sizes. Data pooled from all collections. dler & Franz, 1971). On the other hand, the calyptraeids may feed on detritus stirred up and food particles dropped by the hermit crabs. If this feeding hypothesis is correct, the snails on hermit-crab-occupied-shells are expected to be larger than those on empty shells. No such difference was observed in this study (see Results). Space available on the substratum might be an impor- tant limiting factor (Hendler & Franz, 1971). The sizes of Calyptraea morbida were different on various substra- tum types. This size difference might be explained by the fact that shells of Turritella terebra are larger than those of Turricula javana and Nassarius clathrata. The space limitation hypothesis is supported by the fact that sub- stratum-related difference was found only in females which are larger than males (Table 3). Shell Morphology Shell morphology differs between males and females in shell margin curvature (Figure 1). The shell shape in the family Calyptraeidae is a variable character, strongly affected by the shape of the substratum (Hoagland, 1977a; Vermeij, 1989). It is likely that when a snail fre- quently changes its position, the extent of irregularity in shell margin will be less. Therefore, it is speculated that males, with high RH, might be moving more frequently than females. One might also suspect that the small size itself would render the RH of males higher than that of females. But for snails in the same size range, males were found to have higher RH than females (see Results). This result is compatible with the hypothesis that males may be more mobile than females. Size Structure, Reproductive Characteristics and Sex Change Females were larger than males in Calyptraea morbida (Table 1). This sexual dimorphism in size can be ex- The Veliger, Vol. 43, No. 3 is 2000K5 S rite oS fw = = 2 17504 r“=0.40,n=12 Ha p<0.05 vo 2. n ° > fe 2 E v el iS) fe 2 = | e = 250 = ps Ost T T T T 4 is 100 150 200 250 300 350 S is Body weight (mg) Figure 8 Calyptraea morbida. The relationship between total number of embryos and female body weight. plained by differential mortalities, differential growth rates, or sex change (Wright & Lindberg, 1982). The fol- lowing evidence in the present study is compatible with the hypothesis that individuals of C. morbida changed from male to female. First, the relative frequencies of females increased with size of C. morbida (Figure 2). All small snails (< 18 mg body weight) were males and all larger snails (> 202 mg body weight) were females throughout the year (Table 1). This pattern is compatible with the sex change hypothesis. Second, if this sexual dimorphism in size was caused by differential mortalities or growth rates, we expect to find some small recruited females. Calyptraea morbida recruits year round in Tai- wan (Figures 2, 6). If sex change does not occur in its life history, small recruited females should be found throughout the year. However, all small snails were males in each sample (Table 1). This is incompatible with the differential mortality and the growth rate hypotheses. Third, penis length decreased with increasing size among females (Figure 5) and large females had only a vestigial penis. This phenomenon also occurs in other sex-chang- ing snails in the Calyptraeidae (Fretter & Graham, 1962). Fourth, the overall sex ratio is biased toward males (Table 1) as expected in protandric species (Charnov & Bull, 1989). This phenomenon does not support the differential mortality hypothesis which predicts that the larger, long- lived females should outnumber males. All the above re- sults strongly suggest that sex-change occurs in C. mor- bida, although no gonad of individuals was found with both sperm and oocytes together in the histological study (the same phenomenon also occurred in the protandric Coralliophila violacea [Chen et al., 1998]). The size advantage hypothesis as it is applied to mol- M.-H. Chen & K. Soong, 2000 lusks assumes that high female fecundity is related to large body size but high male reproductive success is re- lated to mobility and therefore often to small size (Hoag- land, 1978). In Calyptraea morbida, \arge females pro- duce more embryos (Figure 8). The relationship between size and reproductive success in males, however, is not clear in this species. Timing of Sex Change The timing of sex change might depend on the size or age of the snails, or environmental factors such as food supply or the presence of conspecific neighbors. The fol- lowing evidence in this study is more compatible with the hypothesis that Calyptraea morbida might change sex at a particular stage. First, if this snail changed sex at a certain size, the size overlap of two sexes should be narrow. How- ever, the overlap in size of the two sexes spanned more than an order of magnitude (18-202 mg) (Table 1). Sec- ond, the “‘recruitment”’ of females, via male sex change, occurred almost year round, but was the highest in April (Figure 4). It is possible that males, with a peak recruit- ment between May and July (Figure 3), changed sex the next year after about 10 months of growth. In the conge- neric C. chinensis, sex change occurs between the first and the second breeding periods when the males are 2—3 years old (Pellegrini, 1949; Bacci, 1951). Acknowledgments. The authors wish to thank Dr. S. K. Wu for critically reading and correcting the manuscript; Dr. C. A. Chen, Mr. Y. W. Chiu, and Mr. Y. S. Shiau for helping with field col- lection; Mr. H. T. Shih for identifying the hermit crabs; Mr. S. L. Wu for drawing the pictures of shells; and Dr. C. M. Lalli and Dr. C. A. Chen for their valuable comments on an early draft of this manuscript. This investigation was partially sponsored by a grant (NSC 85-2321-B-110-001BH) from the National Science Council and a grant from The National Museum of Marine Bi- ology-Aquarium, Taiwan, Republic of China. LITERATURE CITED Baccl, G. 1951. Lermafroditismo di Calyptraea chinensis L. e di altri Calyptraeidae. Pubblicazioni della Stazione Zoolo- gica di Napoli 31:66—90. BERGLUND, A. 1986. Sex change by a polychaete: effects of so- cial and reproductive costs. Ecology 67:837—845. CHARNOV, E. L. 1982. The Theory of Sex Allocation. Princeton University Press: Princeton, New Jersey. 355 pp. CHARNOV, E. L. & J. J. BULL. 1989. Non-Fisherian sex ratios with sex change and environmental sex determination. Na- ture 338:148—150. CHEN, M. H, Y. W. YANG & K. SOONG. 1998. Preliminary ob- servations on change of sex by the coral-inhabiting snails Coralliophila violacea (Lamarck) (Gastropoda: Corallio- philidae). Journal of Experimental Marine Biology and Ecol- ogy 230:207-212.. Cog, W. R. 1932. Sexual phases in the American oyster (Ostrea virginica). Biological Bulletin 63:419—441. Coe, W. R. 1938. Influence of association on the sexual phases Page 217 of gastropods having protandric consecutive sexuality. Bio- logical Bulletin 75:274—285. COLLIN, R. 1995. Sex, size, and position: a test of models pre- dicting size at sex change in the protandrous gastropod Cre- pidula fornicata. American Naturalist 146:815—831. FRETTER, V. & A. GRAHAM. 1962. British Prosobranch Molluscs: Their Functional Anatomy and Ecology. Ray Society: Lon- don. 755 pp. GHISELIN, M. T. 1969. The evolution of hermaphroditism among animals. Quarterly Review of Biology 44:189—208. HELLER, J. 1993. Hermaphroditism in molluscs. Biological Jour- nal of the Linnean Society 48:19—42. HENDLER, G. & D. R. FRANZ. 1971. Population dynamics and life history of Crepidula convexa Say (Gastropoda: Prosobran- chia) in Delaware Bay. Biological Bulletin 141:514—526. HOAGLAND, K. E. 1977a. A gastropoda color polymorphism: one adaptive strategy of phenotypic variation. Biological Bulle- tin 152:360—372. HOAGLAND, K. E. 1977b. Systematic review of fossil and recent Crepidula and discussion of evolution of the Calyptraeidae. Malacologia 16:353—420. HOAGLAND, K. E. 1978. Protandry and the evolution of environ- mentally-mediated sex change: a study of the Mollusca. Ma- lacologia 17:365—391. KARLSON, R. H. & M. A. CarioLtou. 1982. Hermit crab shell colonization by Crepidula convexa Say. Journal of Experi- mental Marine Biology and Ecology 65:1—10. Lessios, H. A. 1992. Testing electrophoretic data for agreement with Hardy-Weinberg expectations. Marine Biology 112: 517-523. PELLEGRINI, O. 1949. Ermafroditismo proterandrico in Calyp- traea chinensis (L.) (Gastropoda Prosobranchiata). Bolletti- no di Zoologia 16:49—59. POLICANSKY, D. 1982. Sex change in plants and animals. Annual Review of Ecology and Systematics 13:471—495. PREMOLI, M. C. & G. SELLA. 1995. Sex economy in benthic poly- chaetes. Ethology Ecology and Evolution 7:27—48. SHENK, M. A. & H. KARLSON. 1986. Colonization of a shell re- source by calyptraeid gastropods: tests of habitat selection and preemption models. Journal of Experimental Marine Bi- ology and Ecology 99:79-89. VERMEU, G. J. 1989. Habitat and form of Crepidula grandis in Japan, with comments on habitat specialization in calyp- traeid gastropods. The Nautilus 103:89-—91. WARNER, R. R. 1975. The adaptive significance of sequential her- maphroditism in animals. American Naturalist 109:61—84. WARNER, R. R. 1988. Sex change and the size-advantage model. Trends in Ecology and Evolution 3:133—136. WARNER, R. R., D. L. Fitcu & J. D. STANDISH. 1996. Social control of sex change in the shelf limpet, Crepidula norri- siarum: size-specific responses to local group composition. Journal of Experimental Marine Biology and Ecology 204: 155-167. WRIGHT, W. G. 1988. Sex change in the Mollusca. Trends in Ecology and Evolution 3:137—140. WRIGHT, W. G. & D. R. LINDBERG. 1982. Direct observation of sex change in the patellacean limpet Lottia gigantea. Journal of Marine Biological Association (U.K.) 62:737—738. Wyatt, H. V. 1960. Protandry and self-fertilization in the Ca- lyptraeidae. Nature 187:520. Wyatt, H. V. 1961. The reproduction, growth and distribution of Calyptraea chinensis (L.). Journal of Animal Ecology 30: 283-302. THE VELIGER © CMS, Inc., 2000 The Veliger 43(3):218—247 (July 3, 2000) Helicarionid Snails of Mounts Mahermana, [lapiry, and Vasiha, Southeastern Madagascar KENNETH C. EMBERTON Florida Museum of Natural History, Box 117800, Gainesville, Florida 32611-7800, USA AND TIMOTHY A. PEARCE! Delaware Museum of Natural History, Box 3937, Wilmington, Delaware 19807-0937, USA Abstract. Quantitative, replicated altitudinal transects yielded 30 helicarionid species in five genera in four subfam- ilies. Descriptions are given of Kalidos balstoni (Angas, 1877); K. fenni sp. nov.; K. prenanti Fischer-Piette, Blanc, Blanc & Salvat, 1994; K. richardi sp. nov.; K. striaspiralis sp. nov.; K. vasihae sp. nov.; K. zahamenensis Fischer-Piette, Blanc, Blanc & Salvat, 1994; Kaliella barrakporensis (Pfeiffer, 1852); Malagarion tillieri sp. nov.; Microcystis andria- mahajai sp. nov.; Mic. basampla sp. nov.; Mic. blanci sp. nov.; Mic. castanea sp. nov.; Mic. compacta sp. nov.; Mic. esetra sp. nov.; Mic. ilapiriensis sp. nov.; Mic. mahermanae sp. nov.; Mic. subangulata sp. nov.; Mic. subplanata sp. nov.; Mic. vohimenae sp. nov.; Mic. vohimenoides sp. nov.; Sitala aliceae sp. nov.; S. amabilis Fischer-Piette & Salvat, 1966; S. elegans sp. nov.; S. euconuliforma sp. nov.; S. gaudens Fischer-Piette & Salvat, 1966; S. ilapiryae sp. nov.; S. josephinae sp. nov.; S. soa sp. nov.; and S. vasihae sp. nov. INTRODUCTION This paper is the final in a series of four that identify and describe the species reported on as morphospecies by Emberton et al. (1996, 1999) and Emberton (1997). This paper treats the Mahermana-Ilapiry-Vasiha helicarionids. MATERIALS AND METHODS Collecting methods have been detailed by Emberton et al. (1996). Sixteen stations were collected and numbered in the “‘Tol’”’ series (for Tolagnaro = Fort Dauphin, the nearest city). These stations have been mapped by Em- berton et al. (1996, 1999) and in Emberton (1997). To shorten the taxonomic descriptions, stations are described briefly below. Catalogued station numbers, given in pa- rentheses, are in the series of the Molluscan Biodiversity Institute (MBI). All stations were restricted to primary forest that had no more than limited selective cutting. Ecological data are given by Emberton (1997:table 1). All stations are in Madagascar: Tulear Province. Mount Mah- ermana (Vohimena Chain) is northeast of the village of Esetra, Nlapiry (Vohimena Chain) is west of Mahialambo, and Vasiha (Anosy Chain) is west of Malio. Latitude and longitude are given in degrees, minutes, and seconds. MBI 373 (= Tol-1). Summit of Mt. Mahermana, 340 m, 24°26'12"S, 47°13'13’E. ‘To whom reprint requests should be sent. MBI 374 (= Tol-2). Slope of Mt. Mahermana, 300 m, 24°26'17"S, 47°13'10"E. MBI 375 (= Tol-3). Slope of Mt. Mahermana, 200 m, 24°26'15"S, 47°13'04’E. MBI 376 (= Tol-4). Valley on Mt. Mahermana, 100 m, 24°26'22"S, 47°12'41"E. MBI 377 (= Tol-5). Summit of Mt. Ilipiry, 540 m, 24°51'40"S, 47°00'20"E. MBI 378 (= Tol-6). Ridge on Mt. lipiry, 500 m, 24°51'33"S, 47°00'27"E. MBI 379 (= Tol-7). Ridge, valley, and slope on Mt. llipiry, 400 m, 24°51'27"S, 47°00'38’E. MBI 380 (= Tol-8). Slope of Mt. Ilipiry, 300 m, 24°51'36"S, 47°00'40"E. MBI 381 (= Tol-9). Slope of Mt. Ilipiry, 200 m, 24°51'39"S, 47°00'46"E. MBI 382 (= Tol-10). Lower summit of Mt. Vasiha, 860 m, 24°55'18"S, 46°44'19"E. MBI 383 (= Tol-11). Slope of Mt. Vasiha, 700 m, 24°55'23"S, 46°44'27'E. MBI 384 (= Tol-12). Slope of Mt. Vasiha, 500 m, 24°55'19"S, 46°44'45"E. MBI 385 (= Tol-13). Valley on Mt. Vasiha, 400 m, 24°55'25"S, 46°44'45"E. MBI 386 (= Tol-14). Slope of Mt. Vasiha, 300 m, 24°55'37"S, 46°44'49"E. MBI 387 (= Tol-15). Slope of Mt. Vasiha, 200 m, 24°56'13"S, 46°45'13"E. MBI 388 (= Tol-16). Slope of Mt. Vasiha, 100 m, 24°56'20"S, 46°46'07"E. K. C. Emberton & T. A. Pearce, 2000 MBI 389 (= Tol-3-4). Incidental collecting between Tol-3 and Tol-4. MBI 390 (= Tol-1-2). Incidental collecting between Tol-1 and Tol-2. MBI 391 (= Tol-sub-5). Incidental collecting below summit of Mt. Ilipiry, Tol-5. MBI 392 (= Tol-7-9). Incidental collecting between Tol-7 and Tol-9. Species identifications and comparisons were made us- ing Fischer-Piette et al. (1994) and Emberton (1994). For each species, the holotype or a representative shell was photographed in apertural, apical, and basal views at 6.4X, 10x, 16x, 25, or 40 magnification, and in api- cal view at 40X magnification (Figures 1—39). Shell char- acters were measured, or measured and calculated, or scored from the photographs or from the shells them- selves. A shell-character matrix was prepared and used to code character-state data into the DELTA system (Dall- witz et al., 1993), which was then used to generate nat- ural-language species descriptions. SYSTEMATICS Higher classification follows Ponder & Lindberg (1997), Nordsieck (1986), and Vaught (1989). Type materials are placed in the United States National Museum, Washing- ton, D.C. (USNM); temporarily in the Molluscan Biodi- versity Institute (MBI), whose collections will revert to the Florida Museum of Natural History, Gainesville; and in the Australian Museum, Sydney (AMS); the Muséum national d’Histoire naturelle, Paris (MNHN, which does not assign catalog numbers to its types); and the Academy of Natural Sciences of Philadelphia (ANSP). For paratype localities, use the MBI catalog number to refer to the station numbers (in parentheses) above. MBI catalog numbers consist of station number, period, species num- ber, D (dry) or A (alcohol-preserved), and when appro- priate H (holotype) or P (paratype) and/or R (represen- tative). Class GASTROPODA Clade HETEROBRANCHIA Clade PULMONATA Order STYLOMMATOPHORA Suborder SIGMURETHRA Infraorder HELICIDA Superfamily HELICARIONOIDEA Family HELICARIONIDAE: Subfamily SESARINAE Genus Kaliella Blanford, 1863 Kaliella barrakporensis (Pfeiffer, 1852) (Figure 39) Kaliella sp. 01, Emberton et al., 1996:210. Emberton, 1997: 1147. Page 219 Representative: MBI 374.02DR, Tol-2 (ad). Other specimens: MBI 373.21D (1 ad), MBI 376.19D (2 ad), MBI 377.23A (1 ad), MBI 379.31D (2 ad, 2 juv), MBI 379.31A (1 ad), MBI 380.23D (1 ad), MBI 381.22D (1 ad, 1 juv; AMS c.203461 [1 ad]), MBI 382.24D (1 ad, 1 juv). Description of representative: Shell Size and Shape. Diameter 3.0 mm; height 2.8 mm. Height-diameter ratio 0.94. Whorls 5.4. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.9. Spire angle 85 degrees. Shell slightly domed. Whorl periphery rounded to slightly angular, pre- sutural ridge present. Suture depth one half whorl from aperture is 1.2% of shell diameter. Umbilicus 3% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 49% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.59. Distance between the columellar and upper peristome insertions 81% of aperture width. Penultimate whorl projects into body whorl, occupying 14% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 95 degrees. Apex, First whorl diameter 0.4 mm. First two whorls diameter 0.8 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell with very fine, regularly spaced, crowded transverse lines. Variation: No conspicuous variation in size or shape. Distribution: Mts. Mahermana, Ilapiry, and Vasiha at 100 to 860 m elevation (this paper), and India, Himalaya, Abyssinia, Commores, Mozambique, South Africa, and widespread on Madagascar (Fischer-Piette et al., 1994). Family HELICARIONIDAE: Subfamily MICROCYSTINAE Genus Microcystis Beck, 1837 Microcystis subplanata Emberton & Pearce, sp. nov. (Figures 1, 2, 3, 4) Microcystis sp. 01, Emberton et al., 1996:210. Emberton, 1997:1146, 1150. Holotype: USNM 860818 (ex MBI 373.09DH, Tol-1, ad). Paratypes: MBI 373.09DP (2 ad, 7 juv; AMS C. 203462 [1 ad]; MNHN [1 ad]; ANSP 400840 [1 ad]), MBI 373.09AP (2 ad, 2 juv), MBI 374.19DP (1 ad, 1 juv), Page 220 The Veliger, Vol. 43, No. 3 Figures 1—5 Figures 1—4. Microcystis subplanata Emberton & Pearce, sp. nov., holotype (Figure 1, four views) and paratypes MBI 373.09DP (Figure 2, three views and spire), MBI 373.09DP (Figure 3, one view), and MBI 374.19DP (Figure 4, one view). Figure 5 (four views). Microcystis castanea Emberton & Pearce, sp. nov., holotype. All scale bars | mm. K. C. Emberton & T. A. Pearce, 2000 MBI 375.17DP (3 juv), MBI 375.17AP (2 juv), MBI 390.03DP (1 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: northeast of village of Esetra: Summit of Mt. Mahermana, 340 m, 24°26'12"S, 47°13'13’E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 7.0 mm; height 3.8 mm. Height-diameter ratio 0.55. Whorls 4.5. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.3. Spire angle 145 degrees. Shell not domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aper- ture is 0.7% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.1% of shell diameter. Umbilicus 0% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 47% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.77. Distance between the columellar and upper peristome insertions 85% of aperture width. Penultimate whorl projects into body whorl, occupying 32% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 65 degrees. Apex. First whorl diameter 1.0 mm. First two whorls diameter 1.8 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; incised lines extremely fine, more than 30 lines between sutures. Variation: The spire can be almost entirely flat, and emp- ty shells sometimes bleach white. The largest specimen (station MBI 373) is 8.2 mm in diameter, with 4.2 whorls. Comparisons: Shape similar to Microcystis platysma Emberton, 1994, but two-thirds the diameter for slightly more whorls and with a narrower aperture. At least 50% larger for the same number of whorls as M. argueyrolli Fischer-Piette, Blanc, Blanc & Salvat, 1994. Distribution: Mt. Mahermana, 200—340 m elevation. Etymology: For its somewhat (L. sub-, under) flat (L. plan-) shell shape. Microcystis castanea Emberton & Pearce, sp. nov. (Figure 5) Microcystis sp. 02, Emberton et al., 1996:210. Emberton, 1997:1147, 1150. Page 221 Holotype: USNM 860819 (ex MBI 373.10DH, Tol-1, ad). Paratypes: MBI 373.10DP (1 ad; AMS C.203463 [1 ad]; MNHN [1 ad]), MBI 374.20DP (1 ad), MBI 375.18DP (1 ad; ANSP 400841 [1 ad]). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: southeast slope of Mt. Vasiha, 700 m, 24°55'23"S, 46°44'27’E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 6.2 mm; height 3.8 mm. Height-diameter ratio 0.61. Whorls 4.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.6. Spire angle 140 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 0.3% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 2.2% of shell diameter. Umbilicus 6% of shell diameter. Shell color red-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 50% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.68. Distance between the columellar and upper peristome insertions 81% of aperture width. Penultimate whorl projects into body whorl, occupying 29% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 95 degrees. Apex. First whorl diameter 1.1 mm. First two whorls diameter 2.1 mm. Embryonic sculpture of interrupted spi- ral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; about 15 fine incised lines between sutures. Variation: The largest shell (station MBI 373) has a di- ameter of 7.8 mm, with 4.8 whorls. Comparisons: Fewer and more rapidly expanding whorls than Microcystis arnali Fischer-Piette, Blanc, Blanc & Salvat, 1994, which lacks the spiral grooves of this spe- cies and is light, not dark, in color. Distribution: Mt. Mahermana, 200—340 m elevation. Etymology: For its chestnut (L. castane-) shell color, dark for the genus. Microcystis compacta Emberton & Pearce, sp. nov. (Figures 6, 7) Microcystis sp. 03, Emberton et al., 1996:210. Emberton, 1997:1146, 1150. Holotype: USNM 860820 (ex MBI 378.04DH, Tol-6, ad). Paratypes: MBI 373.18DP (1 juv), MBI 373.18AP (3 ad), MBI 374.21DP (1 juv), MBI 375.23AP (1 juv), MBI 376.12DP (1 ad, 1 juv), MBI 376.12AP (1 ad), MBI 377.16DP (3 ad, 1 juv), MBI 377.16AP (1 juv), MBI 378.04DP (2 juv; AMS C.203464 [1 ad]; MNHN [1 ad]; ANSP 400842 [1 ad]), MBI 378.04AP (2 ad, 1 juv), MBI 379.24DP (1 ad, 4 juv), MBI 379.24AP (2 juv), MBI 380.17DP (1 ad, 1 juv), MBI 380.17AP (2 ad, 2 juv), MBI 381.17DP (2 juv), MBI 382.17DP (13 juv), MBI 382.17AP (2 ad, 2 juv), MBI 383.11DP (1 ad), MBI 384.15DP (1 juv), MBI 387.10DP (2 juv), MBI 387.10AP (1 juv), MBI 391.03DP (1 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: Ridge on east face of Mt. Ilapiry, 500 m, 24°51'3”S, 47°00'27"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 8.4 mm; height 5.9 mm. Height-diameter ratio 0.71. Whorls 5.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.4. Spire angle 125 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 1.1% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 0.7% of shell diameter. Umbilicus 2% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 53% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.66. Distance between the columellar and upper peristome insertions 85% of aperture width. Penultimate whorl projects into body whorl, occupying 30% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 80 degrees. Apex. First whorl diameter 0.7 mm. First two whorls diameter 1.3 mm. Embryonic sculpture of weak spiral ridges then also with weak growth wrinkles. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth, no sculpture other than weak, irregularly spaced growth wrinkles. Variation: There is some slight variation in darkness of The Veliger, Vol. 43, No. 3 shell coloration, but no conspicuous variation in size or shape. Comparisons: Very similar to Microcystis arnali Fi- scher-Piette, Blanc, Blanc & Salvat, 1994, but conspicu- ously more elevated, with a squarer aperture, lacking any trace of peripheral angulation, and with slightly looser coiling. Distribution: Mts. Mahermana, Ilapiry, and Vasiha, 100— 860 m elevation. Etymology: For its compact shape. Microcystis vohimenae Emberton & Pearce, sp. nov. (Figure 8) Microcystis sp. 04, Emberton et al., 1996:209, 210. Ember- ton, 1997:1146, 1150. Holotype: USNM 860821 (ex MBI 373.11DH, Tol-1, ad). Paratypes: MBI 373.11DP (1 ad, 1 juv; AMS C.203465 [1 ad]; MNHN [1 ad]), MBI 373.11AP (1 ad, 1 juv), MBI 375.19DP (1 juv), MBI 375.19AP (1 ad), MBI 376.24AP (1 ad), MBI 377.17DP (2 ad, 3 juv), MBI 377.17AP (1 juv), MBI 378.24AP (1 ad, 1 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: northeast of village of Esetra: Summit of Mt. Mahermana, 340 m, 24°26'12'S, 47°13'13"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 2.2 mm; height 1.4 mm. Height-diameter ratio 0.66. Whorls 4.1. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 5.2. Spire angle 130 degrees. Shell not domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aper- ture is 0.2% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.0% of shell diameter. Umbilicus 1% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 51% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.71. Distance between the columellar and upper peristome insertions 76% of aperture width. Penultimate whorl projects into body whorl, occupying 30% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 110 degrees. Apex. First whorl diameter 0.4 mm. First two whorls diameter 0.7 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell K. C. Emberton & T. A. Pearce, 2000 Page 223 Figures 6-8 Figures 6, 7. Microcystis compacta Emberton & Pearce, sp. nov., holotype (Figure 6) and paratype MBI 376.12DP (Figure 7). Figure 8. Microcystis vohimenae Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. smooth, no sculpture other than weak, irregularly spaced growth wrinkles. Variation: The largest specimen, from station MBI 373, has a diameter of 2.4 mm, with 4.0 whorls. Comparisons: Similar in shape to but smaller and more tightly coiled than Microcystis ilapiriensis sp. nov., and lacking its basal spiral-line sculpture. Similar in size and coiling tightness to M. vohimenoides sp. nov., but without its wide umbilicus, low aperture, and spiral-line sculpture. Distribution: Vohimena chain (Mts. Mahermana and I[]- apiry), 100 to 540 m elevation. Etymology: For the Vohimena Mountain chain, north of Ft. Dauphin. Microcystis ilapiriensis Emberton & Pearce, sp. nov. (Figure 9) Microcystis sp. 05, Emberton et al., 1996:210. Emberton, 1997:1147. Holotype: USNM 860822 (ex MBI 379.04DH, Tol-7, ad). Paratypes: MBI 379.04DP (1 juv; AMS C.203466 [1 ad]). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: Ridge, val- ley, and slope on southsoutheast face of Mt. lapiry, 400 m, 24°51'27"S, 47.00.38E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 3.5 mm; height 2.2 mm. Height-diameter ratio 0.63. Whorls 4.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 3.8. Spire angle 125 degrees. Shell not domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aper- ture is 0.3% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 2.0% of shell diameter. Umbilicus 4% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 53% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.61. Distance between the columellar and upper peristome insertions 79% of aperture width. Penultimate whorl projects into body whorl, occupying 31% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 90 degrees. Apex. First whorl diameter 0.6 mm. First two whorls diameter 1.1 mm. Embryonic sculpture smooth. The Veliger, Vol. 43, No. 3 Post-Embryonic Shell Sculpture. Post-embryonic shell with very fine granulations. Shell base with fine incised spiral lines. Comparisons: Most similar to Microcystis vohimenae sp. nov., but larger, with looser coiling, and with spiral lines on the base. Distribution: Mt. Ilapiry, 400 m elevation. Etymology: For Mount Iapiry, southern Vohimena Chain. Microcystis vohimenoides Emberton & Pearce, sp. nov. (Figure 10) Microcystis sp. 06, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860823 (ex MBI 380.02DH, Tol-8, ad). Paratypes: MBI 380.02DP (0; AMS C.203467 [1 ad]), MBI 382.18DP (1 ad). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: south slope of Mt. Hapiry, 300 m, 24°51'36"S, 47°00'40"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 2.1 mm; height 1.2 mm. Height-diameter ratio 0.59. Whorls 3.7. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 5.0. Spire angle 140 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 1.2% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 2.6% of shell diameter. Umbilicus 8% of shell diameter. Shell color white. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 50% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.63. Distance between the columellar and upper peristome insertions 75% of aperture width. Penultimate whorl projects into body whorl, occupying 32% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 105 degrees. Apex. First whorl diameter 0.4 mm. First two whorls diameter 0.9 mm. Embryonic sculpture of weak spiral striae. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on K. C. Emberton & T. A. Pearce, 2000 Page 225 Figures 9-11 Figure 9. Microcystis ilapiriensis Emberton & Pearce, sp. nov., holotype. Figure 10. Microcystis vohimenoides Emberton & Pearce, sp. nov., holotype. Figure 11. Microcystis esetra Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. Page 226 upper parts of shell; about 10 fine incised lines between sutures. Variation: No conspicuous variation in size or shape. Comparisons: Similar in its small size and extremely tight coiling to Microcystis vohimenae sp. nov., but with a much lower aperture and wider umbilicus, and with a sculpture of incised spiral lines. Distribution: Mts. Ilapiry and Mt. Vasiha, 300 to 860 m elevation. Etymology: For its resemblance to M. vohimenae sp. nov. Microcystis esetra Emberton & Pearce, sp. nov. (Figure 11) Microcystis sp. 07, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860824 (ex MBI 375.03DH, Tol-3, ad; body preserved as MBI 375.03AH). Paratypes: None. Type Locality: Madagascar: Tulear Province: north of Fort Dauphin: northeast of village of Esetra: west slope of Mt. Mahermana, 200 m, 24°26'15"S, 47°13'04"E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 6.6 mm; height 3.9 mm. Height-diameter ratio 0.59. Whorls 4.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.2. Spire angle 150 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 0.9% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.9% of shell diameter. Umbilicus 5% of shell diameter. Shell color orange-brown. Apex. First whorl diameter 0.9 mm. First two whorls diameter 1.9 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; incised lines extremely fine; more than 30 lines between sutures. Comparisons: Like an enlarged version of Microcystis argueyrolli Fischer-Piette, Blanc, Blanc & Salvat, 1994, which it closely resembles in shape and, apparently, sculpture, but with a diameter of 6.6 mm for four whorls, as opposed to 5 mm for five whorls in M. argueyrolli. Distribution: Mt. Mahermana, 200 m elevation. Etymology: For the village of Esetra. The Veliger, Vol. 43, No. 3 Microcystis andriamahajai Emberton & Pearce, sp. nov. (Figure 12) Microcystis sp. 08, Emberton et al., 1996:210. Emberton, 1997:1148, 1150. Holotype: USNM 860825 (ex MBI 375.04DH, Tol-3, ad; body preserved as MBI 375.04AH). Paratypes: MBI 375.04DP (1 juv), MBI 376.13DP (2 ad, 1 juv; AMS C.203468 [1 ad]; MNHN [1 ad]). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: northeast of village of Esetra: west slope of Mt. Mahermana, 200 m, 24°26'15”S, 47°13'04’E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 3.3 mm; height 2.3 mm. Height-diameter ratio 0.70. Whorls 4.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.0. Spire angle 125 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 1.1% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.2% of shell diameter. Umbilicus 5% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 54% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.60. Distance between the columellar and upper peristome insertions 82% of aperture width. Penultimate whorl projects into body whorl, occupying 33% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 110 degrees. Apex. First whorl diameter 0.6 mm. First two whorls diameter 1.1 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth, no sculpture other than weak, irregularly spaced growth wrinkles. Variation: No conspicuous variation in size or shape. Comparisons: Extremely similar to Microcystis sahavon- drononae Emberton, 1994, but even smaller and more tightly coiled, with 4.8 whorls producing a diameter of 3.3 mm, whereas 4.5 whorls produce 3.7 mm in the latter. Also lacking the spiral sculpture of M. sahavondrononae. Much smaller and more tightly coiled than M. anosiana Fischer-Piette, Blanc, Blanc & Salvat, 1994, and with a straight columellar insertion (angled about 30 degrees in M. anosiana). Distribution: Mt. Mahermana, 100 to 200 m elevation. K. C. Emberton & T. A. Pearce, 2000 Page 227 Figures 12 Figure 12. Microcystis andriamahajai Emberton & Pearce, berton & Pearce, sp. nov., holotype. All scale bars 1 mm. Etymology: For Dr. Benjamin Andriamahaja, National Director of the Ranomafana National Park Project. Microcystis blanci Emberton & Pearce sp. nov. (Figure 13) Microcystis sp. 09, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860826 (ex MBI 380.03DH, Tol-8, ad shell; body included in MBI 380.03AP). and 13 sp. nov., holotype. Figure 13. Microcystis blanci Em- Paratypes: MBI 380.03DP (0; AMS C.203469 [1 ad)]), MBI 380.03AP (4 ad, 1 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: south slope of Mt. Ilapiry, 300 m, 24°51'36"S, 47°00'40”E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 3.8 mm; height 2.4 mm. Height-diameter ratio 0.63. Whorls 3.4. Coiling Page 228 The Veliger, Vol. 43, No. 3 tightness (whorl number divided by natural logarithm of shell diameter) 2.5. Spire angle 135 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 1.2% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.8% of shell diameter. Umbilicus 2% of shell diameter. Shell color pale yellow-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 48% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.91. Distance between the columellar and upper peristome insertions 78% of aperture width. Penultimate whorl projects into body whorl, occupying 27% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 105 degrees. Apex. First whorl diameter 0.7 mm. First two whorls diameter 1.4 mm. Embryonic sculpture smooth then with weak growth wrinkles. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; incised lines extremely fine, more than 30 lines between sutures. Comparisons: Unique for its relatively huge aperture. Similar in shape and fragility to Microcystis charpentieri Fischer-Piette, Blanc, Blanc & Salvat, 1994, but with looser coiling and a much smaller adult size (3.8 mm, 3.4 whorls vs. 5.4 mm, 5 whorls). Distribution: Vohimena chain (Mts. Mahermana and I[]- apiry), 1OO—300 m elevation. Etymology: For Dr. Charles P. Blanc, a researcher of Madagascar’s land snails. Microcystis subangulata Emberton & Pearce, sp. nov. (Figure 14) Microcystis sp. 10, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860827 (ex MBI 383.03DH, Tol-11, ad). Paratypes. MBI 382.19DP (1 ad, 1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: southeast slope of Mt. Vasiha, 700 m, 24°55'23”S, 46°44'27’E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 2.2 mm, height 1.2 mm. Height-diameter ratio 0.54. Whorls 3.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.8. Spire angle 135 degrees. Shell slight- ly domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aperture is 0.8% of shell diameter. Umbilicus 10% of shell diameter. Shell color white. Aperture. Lower peristome angle where it meets pari- etal wall (apertural view) 105 degrees. Apex. First whorl diameter 0.3 mm. First two whorls diameter 0.7 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, uregular growth wrinkles, sculpture on shell base as on upper parts of shell; about 15 fine incised lines between sutures. Variation: The largest specimen is from station MBI 382. It has 4.5 whorls and measures 2.6 mm in diameter. Comparisons: Remarkable for its extremely small size and tight coiling. In these it is matched only by Micro- cystis vohimenoides sp. nov., which has a much more elevated shell (height/diameter 0.7 vs. 0.5). Distribution: Mt. Vasiha, 700 to 860 m elevation. Etymology: For the somewhat (L. sub-, under) angulate (L. angul-) shell periphery. Microcystis mahermanae Emberton & Pearce, sp. nov. (Figure 15) Microcystis sp. 11, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860828 (ex MBI 382. OSDH, Tol-10, ad). Paratypes: None. Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: Lower, south summit of Mt. Vasiha, 860 m, 24°55'18”S, 46°44’9”E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 4.7 mm; height 2.6 mm. Height-diameter ratio 0.55. Whorls 4.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.7. Spire angle 150 degrees. Shell not domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aper- ture is 0.8% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 2.4% of shell diameter. Umbilicus 12% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) K. C. Emberton & T. A. Pearce, 2000 Page 229 Figures 14-16 Figure 14. Microcystis subangulata Emberton & Pearce, sp. nov., holotype. Figure 15. Microcystis mahermanae Emberton & Pearce, sp. nov., holotype. Figure 16. Microcystis basampla Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. Page 230 The Veliger, Vol. 43, No. 3 49% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.76. Distance between the columellar and upper peristome insertions 76% of aperture width. Penultimate whorl projects into body whorl, occupying 30% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 95 degrees. Apex. First whorl diameter 0.9 mm. First two whorls diameter 1.7 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, uregular growth wrinkles, sculpture on shell base as on upper parts of shell; incised lines extremely fine, more than 30 lines between sutures. Comparisons: Resembles Microcystis argueyrolli Fi- scher-Piette, Blanc, Blanc & Salvat, 1994, but with con- siderably looser coiling (2.7 vs. 2.3) and a much larger umbilicus (12% vs. 0% of shell diameter). Distribution: Mt. Vasiha, 860 m elevation. Etymology: For Mount Mahermana, northern Vohimena Chain. Microcystis basampla Emberton & Pearce, sp. nov. (Figure 16) Microcystis sp. 12, Emberton et al., 1996:210. Emberton, 1997:1148. Holotype: USNM 860829 (ex MBI 382.06DH, Tol-10, ad). Paratypes: None. Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: Lower, south summit of Mt. Vasiha, 860 m, 24°55'18"S, 46°44'19"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 3.3 mm; height 2.0 mm. Height-diameter ratio 0.59. Whorls 4.4. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 3.7. Spire angle 130 degrees. Shell not domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aper- ture is 0.8% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 2.2% of shell diameter. Umbilicus 6% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 46% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.75. Distance between the columellar and upper peristome insertions 83% of aperture width. Penultimate whorl projects into body whorl, occupying 32% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 50 degrees. Apex. First whorl diameter 0.6 mm. First two whorls diameter 1.0 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; incised lines extremely fine, more than 30 lines between sutures. Comparisons: Unique for its upper-peripherally flattened whorl. Otherwise its shape is most similar to that of Mi- crocystis arnali Fischer-Piette, Blanc, Blanc & Salvat, 1994, which-is very much larger and more loosely coiled. Distribution: Mt. Vasiha, 860 m elevation. Etymology: For the shell being widest (L. ampli-, spa- cious) near the base (L. bas-). Family HELICARIONIDAE: Subfamily ARIOPHANTINAE Genus Kalidos Gude, 1911 Kalidos balstoni (Angas, 1877) (Figure 17) Kalidos sp. 01, Emberton et al., 1996:210. Emberton, 1997: 1146, 1150. Representative: MBI 373.12DR, Tol-1 (ad). Other specimens: MBI 373.12D (13 juv), MBI 373.12A (7 ad, 1 juv), MBI 374.22D (1 ad, 2 juv), MBI 374.22A (1 ad, 6 juv), MBI 375.20D (1 ad, 8 juv), MBI 375.20A (14 ad, 2 juv), MBI 376.14D (5 juv), MBI 377.18D (7 juv), MBI 377.18A (3 juv), MBI 378.16D (4 juv), MBI 378.16A (2 ad, 4 juv), MBI 379.25D (1 ad, 7 juv), MBI 380.18D (1 ad, 12 juv), MBI 380.18A (6 ad, 4 juv), MBI 381.18D (1 ad, 3 juv; AMS C.203470 [1 ad]; MNHN [1 ad]; ANSP 400843 [1 ad]), MBI 381.18A (1 juv), MBI 382.20D (1 juv), MBI 383.12D (7 juv), MBI 383.12A (3 ad), MBI 384.16D (1 juv), MBI 384.16A (1 ad, 1 juv), MBI 385.13D (1 juv), MBI 389.02A (9 ad, 2 juv), MBI 390.04D (3 juv), MBI 390.04A (1 juv), MBI 392.01A (1 ad). Description of representative: Shell Size and Shape. Diameter 31.6 mm; height 18.9 mm. Height-diameter ratio 0.60. Whorls 5.4. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 1.6. Spire angle 125 degrees. Shell not domed. Whorl periphery angular with or without a keel. Suture depth one half whorl from aperture is 0.5% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; K. C. Emberton & T. A. Pearce, 2000 Page 231 Figures 17 and 18 Figure 17. Kalidos balstoni (Angas, 1877), representative from summit of Mt. Mahermana. Figure 18. Malagarion tillieri Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. The Veliger, Vol. 43, No. 3 width from suture to sub-sutural line (apical view) 0.2% of shell diameter. Umbilicus 5% of shell diameter. Shell color red-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 48% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.75. Distance between the columellar and upper peristome insertions 79% of aperture width. Penultimate whorl projects into body whorl, occupying 17% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 95 degrees. Apex. First whorl diameter 1.7 mm. First two whorls diameter 4.2 mm. Embryonic sculpture of weak spiral ridges and weak transverse ribs. Post-Embryonic Shell Sculpture. Post-embryonic shell with forwardly ascending, closely spaced incised spiral lines cross-cut perpendicularly by incised transverse lines giving the appearance of woven fabric, and with weaker, regularly spaced growth ribs not parallel to either set of incised lines. Variation: No conspicuous variation in size or shape. Distribution: Mts. Mahermana, Ilapiry, and Vasiha at 100 to 860 m elevation (this paper), and widespread in the central, southern, and eastern part of Madagascar (Fi- scher-Piette et al., 1994). Kalidos richardi Emberton & Pearce, sp. nov. (Figures 19, 20) Kalidos sp. 02, Emberton et al., 1996:210. Emberton, 1997: 1146, 1150. Holotype: USNM 860830 (ex MBI 374.03DH, Tol-2, ad). Paratypes: MBI 373.19DP (1 juv), MBI 374.03DP (1 ad, 1 juv; AMS C. 203471 [1 ad]; MNHN [1 ad]; ANSP 400844 [1 ad]), MBI 376.15DP (1 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: northeast of village of Esetra: WSW slope of Mt. Mahermana, 300 m, 24°26'17"S, 47°13'10"E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 11.6 mm; height 7.3 mm. Height-diameter ratio 0.64. Whorls 4.9. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.0. Spire angle 125 degrees. Shell not domed. Whorl periphery angular with or without a keel. Suture depth one half whorl from aperture is 1.4% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 0.2% of shell diameter. Umbilicus 8% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 50% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.70. Distance between the columellar and upper peristome insertions 86% of aperture width. Penultimate whorl projects into body whorl, occupying 23% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 70 degrees. Apex. First whorl diameter 1.2 mm. First two whorls diameter 2.8 mm. Embryonic sculpture of about 30 spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell with very fine, regularly spaced, crowded transverse lines. Variation: The largest shell measures 13.3 mm in di- ameter, with 5.2 whorls (station MBI 374). Comparisons: Nearly identical in size, shape, and coiling tightness to Kalidos prenanti Fischer-Piette, Blanc, Blanc & Salvat, 1994, but with very different embryonic sculp- ture of dense spiral grooves (vs. transversely, then spirally aligned granules) and post-embryonic smooth, featureless sculpture (vs. cross-hatched riblets), and with a weaker peripheral carination. Distribution: Mts. Mahermana and Vasiha, 100—860 m elevation. Etymology: For our guide, collector, and friend, Richard Ialy of Esetra. Kalidos zahamenensis Fischer-Piette, Blanc, Blanc & Salvat, 1994 (Figures 21, 22) Kalidos sp. 03, Emberton et al., 1996:210. Emberton, 1997: 1147, 1150. Representative: MBI 378.05DR, Tol-6 (ad). Other specimens: MBI 376.16D (1 ad, 3 juv; AMS C. 203472 [1 ad]), MBI 379.26D (1 juv), MBI 381.19D (1 juv). Description of representative: Shell Size and Shape. Diameter 9.2 mm; height 4.7 mm. Height-diameter ratio 0.51. Whorls 5.0. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.3. Spire angle 140 degrees. Shell not domed. Whorl periphery angular with or without a keel. Suture depth one half whorl from aperture is 1.1% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.5% K. C. Emberton & T. A. Pearce, 2000 Page 233 Figures 19-22 Figures 19, 20. Kalidos richardi Emberton & Pearce, sp. nov., holotype. Figures 21, 22. Kalidos zahamenensis Fischer-Piette, Blanc, Blanc & Salvat, 1994, representative from Mt. Ilapiry at 500 m. All scale bars | mm. Page 234 of shell diameter. Umbilicus 10% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 47% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.67. Distance between the columellar and upper peristome insertions 81% of aperture width. Penultimate whorl projects into body whorl, occupying 24% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 75 degrees. Apex. First whorl diameter 1.1 mm. First two whorls diameter 2.4 mm. Embryonic sculpture of about 30 spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; about 15 fine incised lines between sutures. Variation: The largest of the specimens reported here has 4.8 whorls, with a diameter of 11.3 mm (station MBI 376). Comparisons: Smaller for the same number of whorls than Kalidos lapillus Fischer-Piette & Bedoucha, 1966; K. anobrachys (Dohrn, 1882); K. prenanti Fischer-Piette, Blanc, Blanc & Salvat, 1994; K. montis Fischer-Piette & Bedoucha, 1966; and K. guernesti Fischer-Piette, Blanc, Blanc & Salvat, 1994. More densely and lightly striate than, and more depressed and less carinate than, K. fallax Fischer-Piette, Blanc & Salvat, 1975. Distribution: Vohimena chain (Mts. Mahermana and I]- apiry) from 100 to 500 m elevation (this paper), and from Zahamena, east-central Madagascar (Fischer-Piette et al., 1994). Kalidos prenanti Fischer-Piette, Blanc, Blanc & Salvat, 1994 (Figures 23, 24) Kalidos sp. 04, Emberton et al., 1996:210. Emberton, 1997: 1147, 1150. Representative: MBI 380.04DR, MBI 380.04AR, Tol-8 (ad). Other specimens: MBI 379.27D (1 ad, 2 juv), MBI 380.04D (1 juv; AMS C.203473 [1 ad]), MBI 380.04A (3 juv), MBI 385.18A (1 juv). Description of representative: Shell Size and Shape. Diameter 10.8 mm; height 6.8 mm. Height-diameter ratio 0.63. Whorls 5.3. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.2. Spire angle 130 degrees. Shell mod- erately domed. Whorl periphery angular with or without The Veliger, Vol. 43, No. 3 a keel. Suture depth one half whorl from aperture is 0.6% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 0.4% of shell diameter. Umbilicus 2% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 48% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.64. Distance between the columellar and upper peristome insertions 82% of aperture width. Penultimate whorl projects into body whorl, occupying 20% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 75 degrees. Apex. First whorl diameter 1.2 mm. First two whorls diameter 2.6 mm. Embryonic sculpture of spiral ridges then also with forward-descending small ribs. Post-Embryonic Shell Sculpture. Post-embryonic shell with forwardly ascending, closely spaced incised spiral lines cross-cut perpendicularly by incised transverse lines giving the appearance of woven fabric, and with weaker, regularly spaced growth ribs not parallel to either set of incised lines. Variation: The largest specimen reported here was from MBI 379: diameter 11.5 mm, whorls 5.4. The single ju- venile specimen from the distant station MBI 385 has similar sculpture, but larger embryonic whorls and a greater rate of whorl expansion; probably it is a different, undescribed species. Distribution: Mt. [lapiry, 300 to 400 m elevation (this paper), and Ambodiriakely forest near Andringitra Re- serve (Fischer-Piette et al., 1994). Comments: With a weak carina more similar to Kalidos torfani Fischer-Piette, Blanc, Blanc & Salvat, 1994, of which this species is probably a synonym. Kalidos fenni Emberton & Pearce, sp. nov. (Figures 25, 26) Kalidos sp. 05, Emberton et al., 1996:210. Emberton, 1997: 1147. Holotype: USNM 860831 (ex MBI 379.05DH, Tol-7, ad). Paratypes: MBI 379.05DP (6 juv; AMS C.203474 [1 ad]), 380.19DP (1 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: Ridge, val- ley, and slope on southsoutheast face of Mt. Ilapiry, 400 m, 24°51'27"S, 47°00'38”E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 12.2 mm; height 7.0 K. C. Emberton & T. A. Pearce, 2000 Page 235 Figures 23—26 Figures 23, 24. Kalidos prenanti Fischer-Piette, Blanc, Blanc & Salvat, 1994, representative from Mt. Ilapiry at 300 m. Figures 25, 26. Kalidos fenni Emberton & Pearce, sp. nov., holotype. All scale bars 1 mm. Page 236 mm. Height-diameter ratio 0.57. Whorls 4.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 1.9. Spire angle 135 degrees. Shell not domed. Whorl periphery angular with or without a keel. Suture depth one half whorl from aperture is 1.7% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 0.3% of shell diameter. Umbilicus 2% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 47% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.76. Distance between the columellar and upper peristome insertions 78% of aperture width. Penultimate whorl projects into body whorl, occupying 25% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 75 degrees. Apex. First whorl diameter 1.2 mm. First two whorls diameter 2.8 mm. Embryonic sculpture of weak spiral ridges then also with weak growth wrinkles. Post-Embryonic Shell Sculpture. Post-embryonic shell with very fine, regularly spaced, crowded transverse lines. Comparisons: Somewhat similar to both Kalidos ber- nardi Fischer-Piette, Blanc, Blanc & Salvat, 1994, and K. helleri Fischer-Piette, Blanc, Blanc & Salvat, 1994, but lacks the distinct body-whorl sculpture of each and has a proportionately smaller aperture than both. Neither of the latter has this species’ diagnostic embryonic sculpture of dense spiral ridges. More loosely coiled and larger than the very similar K. vasihae sp. nov., and lacking its em- bryonic sculpture of beaded spiral ridges. Distribution: Mt. Hapiry, 400 m elevation. Etymology: For Marc Fenn, World Wide Fund for Na- ture, Fort Dauphin. Kalidos striaspiralis Emberton & Pearce, sp. nov. (Figure 27) Kalidos sp. 06, Emberton et al., 1996:210. Emberton, 1997: 1147. Holotype: USNM 860832 (ex MBI 377.04DH, Tol-5, ad). Paratypes: MBI 377.04DP (1 juv), MBI 378.17DP (1 ad, 1 juv), MBI 379.28DP (0; AMS C.203475 [1 ad]), MBI 380.20DP (3 juv), MBI 381.20DP (2 juv). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: Summit of Mt. Ilapiry, 540 m, 24°51'40"S, 47°00'20’E: primary rain- forest. The Veliger, Vol. 43, No. 3 Description of holotype: Shell Size and Shape. Diameter 9.5 mm; height 5.1 mm. Height-diameter ratio 0.54. Whorls 5.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.3. Spire angle 130 degrees. Shell slight- ly domed. Whorl periphery angular with or without a keel. Suture depth one half whorl from aperture is 1.1% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.3% of shell diameter. Umbilicus 10% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 45% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.64. Distance between the columellar and upper peristome insertions 73% of aperture width. Penultimate whorl projects into body whorl, occupying 22% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 105 degrees. Apex. First whorl diameter 1.0 mm. First two whorls diameter 1.9 mm. Embryonic sculpture of beaded spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; about 20 fine incised lines between sutures. Variation: Largest specimen: diameter 12.3 mm, whorls 5.0 (MBI 379). Comparisons: Similar in shape to Kalidos decaryi Fi- scher-Piette, Blanc & Salvat, 1975, and K. propeanob- rachis Fischer-Piette & Bedoucha, 1966, but with much tighter coiling than either. Very distinctive for its sculp- ture of strongly incised spiral grooves and its large um- bilicus. Distribution: Mt. [lapiry, 200 to 540 m elevation. Etymology: For the spiral (L. spiro-) striae (L. striat-) on the shell. Kalidos vasihae Emberton & Pearce, sp. nov. (Figure 28) Kalidos sp. 07, Emberton et al., 1996:210. Emberton, 1997: 1146, 1150. Holotype: USNM 860833 (ex MBI 382.07DH, Tol-10, ad). Paratypes: MBI 382.07DP (1 ad, 6 juv; AMS C.203476 [1 ad]; MNHN [1 ad]), MBI 382.07AP (4 juv), MBI 383.13DP (2 ad, 3 juv; ANSP 400845 [1 ad]), MBI K. C. Emberton & T. A. Pearce, 2000 Page 237 Figures 27 and 28 Figure 27. Kalidos striaspiralis Emberton & Pearce, sp. nov., holotype (three views) and broken paratype MBI 380.20DP (two apical and one side view). Figure 28. Kalidos vasihae Emberton & Pearce, sp. nov., holotype (three views and enlarged side view) and paratype MBI 386.11DP (two apical views). All scale bars | mm. Page 238 385.14DP (2 ad, 1 juv), MBI 385.14AP (1 juv), MBI 386.11DP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: lower, south summit of Mt. Vasiha, 860 m, 24°55'18’S, 46°44'19"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 10.6 mm; height 4.8 mm. Height-diameter ratio 0.46. Whorls 5.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.2. Spire angle 140 degrees. Shell slight- ly domed. Whorl periphery angular with or without a keel. Suture depth one half whorl from aperture is 0.6% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 0.4% of shell diameter. Umbilicus 1% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 46% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.68. Distance between the columellar and upper peristome insertions 87% of aperture width. Penultimate whorl projects into body whorl, occupying 26% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 65 degrees. Apex. First whorl diameter 1.0 mm. First two whorls diameter 2.2 mm. Embryonic sculpture of beaded spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth with extremely fine incised spiral lines and weak, irregular growth wrinkles, sculpture on shell base as on upper parts of shell; incised lines extremely fine, more than 30 lines between sutures. Variation: No conspicuous variation in size or shape. Another specimen from the holotype’s station (MBI 382) is just under it in size: diameter 10.5 mm, whorls 4.7. Comparisons: More tightly coiled and smaller than the very similar K. fenni sp. nov., which lacks this species’ embryonic sculpture of beaded spiral ridges. Distribution: Mt. Vasiha, 400 to 860 m elevation. Etymology: For Mount Vasiha, southern Anosy Moun- tain Chain. Genus Malagarion Tillier, 1979 Malagarion tillieri Emberton & Pearce, sp. nov. (Figure 18) Malagarion sp. 01, Emberton et al., 1996:210. Emberton, 1997:1147, 1150. The Veliger, Vol. 43, No. 3 Holotype: USNM 860834 (ex MBI 386. OSDH, Tol-14, ad). Paratypes: MBI 376.25AP (1 ad), MBI 382.25DP (1 juv), MBI 383.15DP (1 juv; MNHN [1 ad]), MBI 383.15AP (1 juv), MBI 384.17DP (2 juv), MBI 386.05DP (1 juv; AMS C.203477 [1 ad]), MBI 386.05AP (1 ad), MBI 387.12DP (3 juv), MBI 390.07AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: south- southeast slope of Mt. Vasiha, 300 m, 24°55’37’S, 46°44'49"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 9.7 mm; height 5.6 mm. Height-diameter ratio 0.58. Whorls 3.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 1.7. Spire angle 155 degrees. Shell not domed. Whorl periphery rounded to slightly angular, no presutural ridge. Suture depth one half whorl from aper- ture is 0.4% of shell diameter. Sub-sutural line (where inside of shell wall meets previous whorl) visible through translucent shell; width from suture to sub-sutural line (apical view) 1.2% of shell diameter. Shell color pale yel- low-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 53% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.96. Distance between the columellar and upper peristome insertions 149% of aperture width. Penultimate whorl projects into body whorl, occupying 24% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 165 degrees. Apex. First whorl diameter 1.2 mm. First two whorls diameter 1.4 mm. Embryonic sculpture of weak spiral striae. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth, no sculpture other than weak, irregularly spaced growth wrinkles. Variation: The largest specimen (diameter 10.4 mm) is from station MBI 386. Comparisons: Shell shape very similar to Chlamydarion (?) grili Fischer-Piette, Blanc, Blanc & Salvat, 1994, but lacks fine spiral striae of that species and is smaller for the same whorl number. Similar in size and shape to Mal- agarion antalahae Emberton, 1994, but lacks sculpture of that species. Lacks the shallow spiral groove in the upper periphery of Malagarion andampibei Emberton, 1994, and is smaller at the same whorl number. Upper whorls not whitish as in Malagarion andranomenae Em- berton, 1994, and is smaller at the same whorl number, mantle covers less of the shell, and lacks warty protu- berances of that species. Final whorl not ascending on K. C. Emberton & T. A. Pearce, 2000 previous whorl as in Chlamydarion (?) puitsi Fischer- Piette, Blanc, Blanc & Salvat, 1994. Distribution: Mts. Mahermana and Mt. Vasiha, 100—860 m elevation. Etymology: For Dr. Simon Tillier, Musée Nationale d’histoire Naturelle, Paris, who first described and named the genus. Family HELICARIONIDAE: Subfamily MACROCHLAMYDINAE Genus Sitala H. Adams, 1865 Sitala josephinae Emberton & Pearce, sp. nov. (Figure 29) Sitala sp. 01, Emberton et al., 1996:210. Emberton, 1997: 1146, 1150. Holotype: USNM 860835 (ex MBI 375.21DH, Tol-3, ad). Paratypes: MBI 373.20DP (7 juv; AMS C.203478 [1 ad]; MNHN [1 ad]), MBI 373.20AP (1 juv), MBI 374.23DP (3 juv), MBI 374.23AP (1 ad), MBI 375.21DP (1 ad, 1 juv; ANSP 400846 [1 ad]), MBI 375.21AP (2 ad), MBI 376.17DP (1 ad). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: northeast of village of Esetra: west slope of Mt. Mahermana, 200 m, 24°26'15”S, 47°13'04"E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 4.5 mm; height 4.6 mm. Height-diameter ratio 1.03. Whorls 6.3. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.2. Spire angle 80 degrees. Shell slightly domed. Whorl periphery rounded to slightly angular, pre- sutural ridge present. Suture depth one half whorl from aperture is 2.9% of shell diameter. Umbilicus 6% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 48% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.62. Distance between the columellar and upper peristome insertions 82% of aperture width. Penultimate whorl projects into body whorl, occupying 7% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 100 degrees. Apex. First whorl diameter 0.5 mm. First two whorls diameter 1.0 mm. Embryonic sculpture of three spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell with three carinae between sutures; with very weak spiral Page 239 ridges between the carinae, about 10 spiral ridges from the upper suture to the first carina, about seven spiral ridges to the next carina, and about five spiral ridges to the third. Shell base smooth. Variation: No conspicuous variation in size or shape. The largest specimen (station MBI 373) has 6.8 whorls and a height of 5.3 mm. Comparisons: Most similar to Sitala brancsiki Boettger, 1892, in shape and sculpture, both having numerous weaker spiral ridges between stronger spiral ridges, but S. brancsiki has four spiral ridges instead of three, and its ridges are stronger. Distribution: Mt. Mahermana, 100 to 340 m elevation. Etymology: For Josephine Djaohasara Emberton, wife of KCE. Sitala elegans Emberton & Pearce, sp. nov. (Figure 30) Sitala sp. 02, Emberton et al., 1996:210. Emberton, 1997: 1148, 1150. Holotype: USNM 860836 (ex MBI 379.06DH, Tol-7, ad). Paratypes: MBI 379.06DP (2 juv; AMS C.203479 [1 ad]), MBI 379.06AP (4 ad). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: Ridge, val- ley, and slope on southsoutheast face of Mt. Ilapiry, 400 m, 24°51'27"S, 47°00'38"E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 4.2 mm; height 5.2 mm. Height-diameter ratio 1.24. Whorls 6.6. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.6. Spire angle 65 degrees. Shell slightly domed. Whorl periphery rounded to slightly angular, pre- sutural ridge present. Suture depth one half whorl from aperture is 2.4% of shell diameter. Umbilicus 1% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 47% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.68. Distance between the columellar and upper peristome insertions 87% of aperture width. Penultimate whorl projects into body whorl, occupying 12% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 105 degrees. Apex. First whorl diameter 0.8 mm. First two whorls diameter 1.3 mm. Embryonic sculpture of three inter- rupted spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell Pa o 240 The Veliger, Vol. 43, No. 3 Figures 29 and 30 Figure 29, Sitala josephinae Emberton & Pearce, sp. nov., holotype. Figure 30 sp. nov., holotype. All scale bars | mm. . Sitala elegans Emberton & Pearce, K. C. Emberton & T. A. Pearce, 2000 with three carinae between sutures; bristles present on all three carinae, about five bristles per carina in one-tenth of penultimate whorl; bristles about 0.8 mm long. Base of shell smooth. Comparisons: Unique within the genus for its bristly sculpture. Distribution: Mt. [lapiry, 400 m elevation. Etymology: For the elegant (L. elegan-) sculpture and shape. Sitala ilapiryae Emberton & Pearce, sp. nov. (Figure 31) Sitala sp. 03, Emberton et al., 1996:210. Emberton, 1997: 1148, 1150. Holotype: USNM 860837 (ex MBI 379.07DH, Tol-7, ad). Paratypes: MBI 379.07DP (1 ad, 4 juv; AMS C.203480 [1 ad]). Type locality: Madagascar: Tulear Province: north of Fort Dauphin: west of village of Mahialambo: Ridge, val- ley, and slope on southsoutheast face of Mt. Ilapiry, 400 m, 24°51'27”S, 47°00'38’E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 2.5 mm; height 2.0 mm. Height-diameter ratio 0.82. Whorls 4.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.6. Spire angle 85 degrees. Shell slightly domed. Whorl periphery rounded to slightly angular, pre- sutural ridge present. Suture depth one half whorl from aperture is 2.4% of shell diameter. Umbilicus 4% of shell diameter. Shell color white. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 46% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.72. Distance between the columellar and upper peristome insertions 78% of aperture width. Penultimate whorl projects into body whorl, occupying 19% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 100 degrees. Apex. First whorl diameter 0.6 mm. First two whorls diameter 1.1 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell smooth, no sculpture other than weak, irregularly spaced growth wrinkles. Variation: No conspicuous variation in size or shape. Comparisons: More squat than Sitala amabilis Fischer- Piette & Salvat, 1966, and S. ankazobei Fischer-Piette, Blanc, Blanc & Salvat, 1994, and lacking the spiral striae of the latter. Page 241 Distribution: Mt. [lapiry, 400 m elevation. Etymology: For Mount Ilapiry, southern Vohimena chain. Sitala euconuliforma Emberton & Pearce, sp. nov. (Figure 32) Sitala sp. 04, Emberton et al., 1996:210. Emberton, 1997: 1148. Holotype: USNM 860838 (ex MBI 384.04DH, Tol-12, ad). Paratypes: MBI 382.21DP (1 ad, 1 juv; MNHN [1 ad]), MBI 385.15DP (1 ad, 1 juv; AMS C.203481 [1 ad]), MBI 386.12DP (3 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: east slope of Mt. Vasiha, 500 m, 24°55'19"S, 46°44'45”E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 3.4 mm; height 4.1 mm. Height-diameter ratio 1.18. Whorls 7.3. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 6.0. Spire angle 60 degrees. Shell mod- erately domed. Whorl periphery rounded to slightly an- gular, presutural ridge present. Suture depth one half whorl from aperture is 1.5% of shell diameter. Umbilicus 4% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 45% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.69. Distance between the columellar and upper peristome insertions 94% of aperture width. Penultimate whorl projects into body whorl, occupying 17% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 100 degrees. Apex. First whorl diameter 0.7 mm. First two whorls diameter 1.1 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell with very fine granulations. Periphery with a single strong spiral ridge, appearing in the suture as a thread. Variation: No conspicuous variation in size or shape. Comparisons: Has the sutural thread, lack of other spiral sculpture, and similar shape and whorl number as Sitala filomarginata Boettger, 1892, but this is smaller, a little less squat, and the carina is not wavy. Distribution: Mt. Vasiha, 300 to 860 m elevation. Etymology: For the shell shape (L. forma) resembling some North American Euconulus Reinhardt, 1883. Page 242 The Veliger, Vol. 43, No. 3 Figures 31-33 Figure 31. Sitala ilapiryae Emberton & Pearce, sp. nov., holotype. Figure 32. Sitala euconuliforma Emberton & Pearce, sp. nov., holotype. Figure 33 Sitala amabilis Fischer-Piette & Salvat, 1966, representative from Mt. Vasiha at 500 m. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 Sitala gaudens Fischer-Piette & Salvat, 1966 (Figures 34, 35) Sitala sp. 05, Emberton et al., 1996:210, 211. Emberton, 1997:1146, 1150. Representative: MBI 378.06DR, Tol-6 (ad). Other specimens: MBI 376.18D (1 juv), MBI 377.19D (2 ad, 20 juv; AMS C. 203482 [1 ad]; MNHN [1 ad]), MBI 377.19A (1 ad), MBI 378.06D (3 ad, 30 juv; ANSP 400847 [1 ad]), MBI 379.29D (1 ad, 3 juv), MBI 380.21D (1 juy). Description of representative: Shell Size and Shape. Diameter 7.2 mm; height 7.2 mm. Height-diameter ratio 0.99. Whorls 5.9. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 3.0. Spire angle 80 degrees. Shell not domed. Whorl periphery rounded to slightly angular, pre- sutural ridge present. Suture depth one half whorl from aperture is 1.2% of shell diameter. Umbilicus 1% of shell diameter. Shell color pale yellow-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 53% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.74. Distance between the columellar and upper peristome insertions 84% of aperture width. Penultimate whorl projects into body whorl, occupying 19% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 105 degrees. Apex. First whorl diameter 1.0 mm. First two whorls diameter 1.8 mm. Embryonic sculpture smooth. Post-Embryonic Shell Scuplture. Post-embryonic shell with fine, closely spaced, wavy, beaded spiral sculpture, about 40 spiral sculptures between sutures, sculpture con- tinuing on shell base. Variation: The largest specimen reported here (station MBI 377) has 6.2 whorls, height 9.1 mm, diameter 8.8 mm. Distribution: Vohimena chain (Mts. Mahermana and II- apiry), 100-540 m elevation (this paper), and Manjaka- tompo forest in central Madagascar (Fischer-Piette et al., 1994). Comments: The illustration of Sitala gaudens Fischer- Piette & Salvat, 1966, in Fischer-Piette et al. (1994:plate 43:7) is a shell having a more concave spire profile, and the description indicates that the spiral incised lines are weaker than the transverse lines (the opposite is true for this specimen). Page 243 Sitala amabilis Fischer-Piette & Salvat in Fischer- Piette, Bedoucha & Salvat, 1966 (Figure 33) Sitala sp. 06, Emberton et al., 1996:210. Emberton, 1997: 1146. Representative: MBI 384.05DR, Tol-12 (ad). Other specimens: MBI 382.22D (1 juv), MBI 384.05D (3 juv), MBI 388.10D (1 juv). Description of representative: Shell Size and Shape. Diameter 5.2 mm; height 5.4 mm. Height-diameter ratio 1.03. Whorls 5.2. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 3.2. Spire angle 85 degrees. Shell not domed. Whorl periphery rounded to slightly angular, pre- sutural ridge present. Suture depth one half whorl from aperture is 1.9% of shell diameter. Umbilicus 3% of shell diameter. Shell color pale yellow-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 57% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.67. Distance between the columellar and upper peristome insertions 80% of aperture width. Penultimate whorl projects into body whorl, occupying 15% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 120 degrees. Apex. First whorl diameter 0.8 mm. First two whorls diameter 1.5 mm. Embryonic sculpture smooth. Post-Embryonic Shell Sculpture. Post-embryonic shell with very fine granulations. Sculpture on shell base as on upper parts of shell. Distribution: Mt. Vasiha, from 100 to 500 m elevation (this paper); Mandraka south to Andringitra (Fischer-Piet- te et al., 1994). Comments: This is very likely a new species because of its sculptural differences from described Sitala amabilis, but because of its great geographical distance it could be a local variant. Sitala aliceae Emberton & Pearce, sp. nov. (Figure 36) Sitala sp. 07, Emberton et al., 1996:209, 210. Emberton, 1997:1146, 1150. Holotype: USNM 860839 (ex MBI 384.06DH, Tol-12, ad). Paratypes: MBI 377.20DP (7 ad, 3 juv), MBI 377.20AP (2 ad), MBI 378.18DP (7 ad, 1 juv), MBI 378.AP (10 ad, 1 juv), MBI 379.30DP (12 ad, 11 juv), MBI 379.AP (7 ad), MBI 380.22DP (5 ad, 6 juv), MBI 380.22AP (3 ad), Page 244 The Veliger, Vol. 43, No. 3 Figures 34—36 Figures 34, 35. Sitala gaudens Fischer-Piette & Salvat, 1966, representative from Mt. Ilapiry at 500 m. Figure 36. Sitala aliceae Emberton & Pearce, sp. nov., paratype MBI 379.30DP (large shell in one standard view) and holotype (smaller, all other views). All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 MBI 381.21DP (5 ad, 4 juv), MBI 382.23DP (11 ad, 17 juv), MBI 382.23AP (2 ad, 2 juv), MBI 383.14DP (5 ad, 3 juv), MBI 384.06DP (9 ad, 8 juv), MBI 384.06AP (2 juv), MBI 385.16DP (3 ad, 32 juv; AMS C.203483 [1 ad]; MNHN [1 ad]; ANSP 400848 [1 ad]), MBI 385.16AP (6 ad, 13 juv), MBI 386.13DP (12 ad, 21 juv), MBI 386.13AP (2 ad, 6 juv), MBI 387.11DP (6 ad, 15 juv), MBI 387.11AP (3 juv), MBI 388.11DP (5 ad, 2 juv), MBI 388.11AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: east slope of Mt. Vasiha, 500 m, 24°55'19"S, 46°44'45°E: primary rainforest. Description of holotype: Shell Size and Shape. Diameter 2.8 mm; height 2.0 mm. Height-diameter ratio 0.73. Whorls 4.8. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 4.7. Spire angle 110 degrees. Shell mod- erately domed. Whorl periphery rounded to slightly an- gular, presutural ridge present. Suture depth one half whorl from aperture is 2.3% of shell diameter. Umbilicus 7% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 48% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.61. Distance between the columellar and upper peristome insertions 85% of aperture width. Penultimate whorl projects into body whorl, occupying 20% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 90 degrees. Apex. First whorl diameter 0.6 mm. First two whorls diameter 1.0 mm. Embryonic sculpture smooth then with spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell with fine spiral ridges and with weak irregular growth wrinkles; six spiral ridges between sutures. Shell base smooth. Variation: Size and shape vary enormously; the extremes are illustrated in Figure 36. Nearly the full range between these extremes occurs within a single presumed popula- tion (station MBI 386). Comparisons: Unique in its spiral sculpture that is coarse and strong above, fine and weak below. Distribution: Mts. [lapiry and Vasiha, 100 to 860 m el- evation. Etymology: For Alice Doolittle, partner of TAP. Sitala vasihae Emberton & Pearce, sp. nov. (Figure 37) Sitala sp. 08, Emberton et al., 1996:210. Emberton, 1997: 1148. Page 245 Holotype: USNM 860840 (ex MBI 384.07DH, Tol-12, ad). Paratypes: None. Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: east slope of Mt. Vasiha, 500 m, 24°55’19"S, 46°44’45”E: primary rainforest. Description of holotype: Shell Size and Shape. Whorls 4.0. Spire angle 80 de- grees. Shell not domed. Whorl periphery rounded to slightly angular, presutural ridge present. Shell color white. Apex. First whorl diameter 0.5 mm. First two whorls diameter 0.9 mm. Embryonic sculpture of weak spiral ridges then also with weak growth wrinkles. Post-Embryonic Shell Sculpture. Post-embryonic shell with fine spiral ridges and with weak irregular growth wrinkles; about 10 spiral ridges between sutures, no spiral ridges below shell periphery. Comparisons: With fine spiral striae and similar spire shape as a juvenile of Sitala hestia (Dohrn, 1882), but is smaller. Distribution: Mt. Vasiha, 500 m elevation. Etymology: For Mount Vasiha, southern Anosy chain. Sitala soa Emberton & Pearce, sp. nov. (Figure 38) Sitala sp. 09, Emberton et al., 1996:210. Emberton, 1997: 1148. Holotype: USNM 860841 (ex MBI 383.04DH, Tol-11, ad; body preserved as MBI 383.04AH). Paratypes: None. Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: southeast slope of Mt. Vasiha, 700 m, 24°55'23"S, 46°44'27"E: pri- mary rainforest. Description of holotype: Shell Size and Shape. Diameter 6.2 mm; height 3.9 mm. Height-diameter ratio 0.63. Whorls 5.3. Coiling tightness (whorl number divided by natural logarithm of shell diameter) 2.9. Spire angle 130 degrees. Shell mod- erately domed. Whorl periphery rounded to slightly an- gular, presutural ridge present. Suture depth one half whorl from aperture is 0.7% of shell diameter. Umbilicus 10% of shell diameter. Shell color orange-brown. Aperture. Aperture width (measured parallel to a line between the columellar and upper peristome insertions) 47% of shell diameter. Aperture height-width ratio (height measured to and perpendicular to a line between the col- umellar and upper peristome insertions) 0.68. Distance Page 246 The Veliger, Vol. 43, No. 3 Figures 37-39 Figure 37. Sitala vasihae Emberton & Pearce, sp. nov., holotype. Figure 38. Sitala soa Emberton & Pearce, sp. nov., holotype. Figure 39. Kaliella barrakporensis (Pfeiffer, 1852), representative from Mt. Mahermana at 300 m. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 between the columellar and upper peristome insertions 85% of aperture width. Penultimate whorl projects into body whorl, occupying 29% of aperture height measure. Lower peristome angle where it meets parietal wall (ap- ertural view) 65 degrees. Apex. First whorl diameter 0.9 mm. First two whorls diameter 1.7 mm. Embryonic sculpture smooth then with spiral ridges. Post-Embryonic Shell Sculpture. Post-embryonic shell with fine spiral ridges and with weak irregular growth wrinkles; about five weak spiral ridges between sutures. Spiral ridges present below shell periphery but shell base smooth. Comparisons: Unique in its combination of spiral sculp- ture; low, domed shape; and conspicuous umbilicus. Distribution: Mt. Vasiha, 700 m elevation. Etymology: For the beautiful (Malagasy soa) shell. DISCUSSION These descriptions of 30 helicarionids complete the sys- tematic documentation for our previous distributional and ecological analyses of Mahermana-lIlapiry-Vasiha land snails (Emberton et al., 1996, 1999; Emberton, 1997). Acknowledgments. We are grateful to the U.S. National Science Foundation and USAID for funding (grant DEB-9201060 to KCE); to staffs of the Ranomafana National Park Project, the Madagascar Départment des Eaux et Foréts, and the Tolagnaro (Fort Dauphin) office of the World Wide Fund for Nature for logistical aid; to Roger Randalana and assistants from Esetra, Mahialambo, and Malio for collecting; to Felix Rakotomalala for Page 247 curatorial assistance; and to Lucia Emberton for help in mounting the photographs. LITERATURE CITED DatiLwitz, M. J., T. A. PAINE & E. J. ZURCHER. 1993. DELTA User’s Guide: A General System for Processing Taxonomic Descriptions. 4th ed. CSIRO Information Services: Mel- bourne, Australia 136 pp. EMBERTON, K. C. 1994. Thirty new species of Madagascan land snails. Proceedings of the Academy of Natural Sciences of Philadelphia 145:147—189. EMBERTON, K. C. 1997. Diversities, distributions, and abundances of 80 species of minute-sized land snails in southeastern- most Madagascan rainforests, with a report that lowlands are richer than highlands in endemic and rare species. Biodi- versity and Conservation 6:1137—1154. EMBERTON, K. C., T. A. PEARCE & R. RANDALANA. 1996. Quan- titatively sampling land-snail species richness in Madagascar rainforests. Malacologia 38:203—212. EMBERTON, K. C., T. A. PEARCE & R. RANDALANA. 1999. Mol- luscan diversity in the conserved Vohimena and the uncon- served Anosy mountain chains, southeast Madagascar. Bio- logical Conservation 89:183-188. FISCHER-PIETTE, E., C. P. BLANC, EK BLANC & FE Satvart. 1994. Gastéropodes terrestres pulmonés. Faune de Madagascar, 83: 1-552. Norpsieck, H. 1986. The system of the Stylommatophora (Gas- tropoda), with special regard to the systematic position of the Clausiliidae, II. Importance of the shell and distribution. Archiv fiir Molluskenkunde 117:93-116. PONDER, W. E & D. R. LINDBERG. 1997. Towards a phylogeny of gastropod molluscs: an analysis using morphological characters. Zoological Journal of the Linnean Society 119: 83-265. VAUGHT, K. C. 1989. A Classification of the Living Mollusca. American Malacologists, Inc.: Melbourne, Florida. 189 pp. THE VELIGER © CMS, Inc., 2000 The Veliger 43(3):248-264 (July 3, 2000) Charopid Snails of Mounts Mahermana, Hapiry, and Vasiha, Southeastern Madagascar, with Description of a New Genus and with Conservation Statuses of Nine Species KENNETH C. EMBERTON Florida Museum of Natural History, Box 117800, Gainesville, Florida 32611-7800, USA AND TIMOTHY A. PEARCE! Delaware Museum of Natural History, Box 3937, Wilmington, Delaware 19807-0937, USA Abstract. Quantitative, replicated altitudinal transects yielded nine charopids, all belonging to the endemic genus Reticulapex gen. nov., which is characterized by cross-hatched embryonic sculpture; an extremely elongate fertilization pouch-seminal receptacle complex; nearly basal entry of the vas deferens into, and nearly apical origin of the penial- retractor muscle from, the epiphallus; and deposition of a spiral spermatophore on the mate’s penis. Reticulapex gen. nov. represents the fifth known independent evolution of external sperm exchange within the Sty- lommatophora. At least some other charopids have internal sperm exchange. Descriptions are given of Reticulapex gen. nov. apexfortis sp. nov., R. compactus sp. nov., R. fischerpiettei sp. nov., R. flammulatus sp. nov., R. lucidus sp. nov., R. scaber sp. nov., R. subangulatus sp. nov., R. ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., and R. villosus sp. nov. Description of Reticulapex gen. nov. includes the additional new combinations R. choutardi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. druggi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. harananae (Emberton, 1994) comb. nov., R. intridi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. talatai (Emberton, 1994) comb. nov., R. vatuvavyae (Emberton, 1994) comb. nov., and R. vineti (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov. Distributional data allowed evaluation of each of the nine Mahermana-Ilapiry-Vasiha charopid species for its conser- vation status, applying the latest IUCN criteria. Two species are proposed as Critically Endangered and seven as En- dangered. INTRODUCTION (MBI). All stations were restricted to primary forest that had no more than limited selective cutting. Ecological data are given by Emberton (1997:table 1). All stations are in Madagascar: Tulear Province. Mount Mahermana (Vohimena Chain) is northeast of the village of Esetra, Ilapiry (Vohimena Chain) is west of Mahialambo, and Vasiha (Anosy Chain) is west of Malio. Latitude and lon- gitude are given in degrees, minutes, and seconds. MATERIALS anb METHODS Tol-1 (= MBI 373). Summit of Mt. Mahermana, 340 m, 24°26'12”S, 47°13'13”E. Tol-1-2 (= MBI 390). Incidental collecting between This paper is the third in a series of four that identify and describe the species reported on as morphospecies by Emberton et al. (1996, 1999) and Emberton (1997). This paper treats the Mahermana-Ilapiry-Vasiha charopids and evaluates each charopid species for conservation status. Collecting methods have been detailed by Emberton et al. (1996). Sixteen stations were collected and numbered in the “‘Tol’”’ series (for Tolagnaro = Fort Dauphin, the Tol-1 and Tol-2. nearest city). These stations have been mapped by Em- Bere me seo Slope of Mt. Mahermana, 300 m, berton et al. (1996) and by Emberton (1997). To shorten 24°26'17"S, 47°13’ 10°E. the taxonomic descriptions, stations are described briefly lols Ss se Te) Slope of Mt. Mahermana, 200 m, below. Catalogued station numbers, given in parentheses, 24°26°15"S, 47° 13'04'E. : ! are in the series of the Molluscan Biodiversity Institute Tol-3-4 (= MBI 389). Incidental collecting between Tol-3 and Tol-4. Tol-4 (= MBI 376). Valley on Mt. Mahermana, 100 'To whom reprint requests should be sent. m, 24°26'22"S, 47°12'41’E. K. C. Emberton & T. A. Pearce, 2000 Tol-5 (= MBI 377). Summit of Mt. [apiry, 540 m, 24°51'40"S, 47°00'20"E. Tol-sub-5 (= MBI 391). Incidental collecting below summit of Mt. Ilapiry, Tol-5. Tol-6 (= MBI 378). Ridge on Mt. [apiry, 500 m, 24°51'33”"S, 47°00'27’E. Tol-7 (= MBI 379). Ridge, valley, and slope on Mt. Ilapiry, 400 m, 24°51'27"S, 47°00'38’E. Tol-8 (= MBI 380). Slope of Mt. Ilapiry, 300 m, 24°51'36"S, 47°00'40"E. Tol-7-9 (= MBI 392). Incidental collecting between Tol-7 and Tol-9. Tol-9 (= MBI 381). Slope of Mt. Ilapiry, 200 m, 24°51'39"S, 47°00'46’E. Tol-10 (= MBI 382). Lower summit of Mt. Vasiha, 860 m, 24°55'18”"S, 46°44’19”E. Tol-11 (= MBI 383). Slope of Mt. Vasiha, 700 m, 24°55'23"S, 46°44'27"E. Tol-12 (= MBI 384). Slope of Mt. Vasiha, 500 m, 24°55'19"S, 46°44'45"E. Tol-13 (= MBI 385). Valley on Mt. Vasiha, 400 m, 24°55'25"S, 46°44'45"E. Tol-14 (= MBI 386). Slope of Mt. Vasiha, 300 m, 24°55'37"S, 46°44'49"E. Tol-15 (= MBI 387). Slope of Mt. Vasiha, 200 m, 24°56'13"S, 46°45'13”E. Tol-16 (= MBI 388). Slope of Mt. Vasiha, 100 m, 24°56'20"S, 46°46'07’E. Species identifications and comparisons were made us- ing Fischer-Piette et al. (1994) and Emberton (1994a). For each species, the holotype or a representative shell was photographed in apertural, basal, and side views at either 6.4X, 10x, 16x, or 25X magnification, and in apical view at 40 magnification. Additional specimens were photographed as needed to illustrate shell variation or sculptural features. Twenty-four shell characters (Table 1; Emberton & Pearce, 1999:fig. 1, 2000:fig. 1) were measured, or mea- sured and calculated, or scored from the photographs or from the shells themselves. An adult anatomy was available for each of five spe- cies. From each, the reproductive system was removed and photographed as it was turned and progressively dis- sected to expose characters. Thirteen reproductive-ana- tomical characters (Table 1, Figure 12) were taken from the photographs or from the dissections themselves. Character matrices were prepared and used to code character-state data into the DELTA system (Dallwitz et al., 1993), which was then used to generate natural-lan- guage species descriptions. For each charopid species, conservation status was evaluated using the new categories and criteria of the In- ternational Union for the Conservation of Nature (IUCN, 1996). Ranges were estimated from 1992-1995 distribu- tional data (Emberton, unpublished). Rainforest extent and decline were assessed using Green & Sussman Page 249 Table 1 Shell and reproductive characters used in descriptions. Many are illustrated by Emberton & Pearce (1999a:figure 1, 1999b:figure 1). SHELL 1. Diameter (0.1 mm) 2. Height (0.1 mm) 3. Height/Diameter (0.1) 4. Spire angle (degrees) 5. Whorl periphery shape (round, angular, keeled) 6. Whorl shoulder shape (round, flat) 7. Aperture width parallel to parietal callus (% diameter) 8. Aperture height (perpendicular to parietal callus)/width (0.01) 9. Umbilicus size (% diameter) 10. Whorl number (0.1) 11. Coiling tightness (whorl number/In diameter) 12. Embryonic whorl number (0.1) 13. Embryonic shell diameter (0.1 mm) 14. First whorl diameter (0.1 mm) 15. Embryonic sculpture 16. Post-embryonic cross-hatch sculpture (absent, faint, mod- erate, strong) 17. Transverse rib density (number in first 0.1 of body whorl) 18. Rib height (% shell diameter) 19. Rib periostracum locally extended into hair (no, length [% shell diameter]) 20. Periostracal hairs unassociated with ribs (number between sutures) 21. Height of periostracal hairs unassociated with ribs (% shell diameter) 22. Conspicuous spiral grooves below periphery of body whorl (no, number) 23. Shell ground color 24. White blotches of color on shell REPRODUCTIVE SYSTEM 25. Penis length (0.1 mm) 26. Penis length/shell diameter (0.1) 27. Penis width (range, 0.1 mm) 28. Penial sculpture 29. Epiphallus shape and orientation 30. Spermatophore size, shape, and deposition site 31. Penial retractor muscle attachment position 32. Vas deferens entry position and width 33. Atrium length 34. Vagina length 35. Spermathecal duct width and length 36. Spermatheca shape and position 37. Fertilization pouch-seminal receptacle complex shape and length (1990), Sussman et al. (1994), and the most recently available topographic maps. SYSTEMATICS Higher classification follows Ponder & Lindberg (1997) and Nordsieck (1996). Type materials are placed in the United States National Museum, Washington, D.C. Page 250 (USNM); temporarily in the Molluscan Biodiversity In- stitute (MBI), all of whose collections will revert in the near future to the Florida Museum of Natural History, Gainesville; and in the Australian Museum, Sydney (AMS); the Muséum national d’ Histoire naturelle, Paris (MNHN); and the Academy of Natural Sciences of Phil- adelphia (ANSP). For paratype localities, use the MBI catalog number to refer to the station numbers (in paren- theses) above. MBI catalog numbers consist of station number, period, species number, D (dry) or A (alcohol- preserved), and when appropriate H (holotype) or P (par- atype) or R (representative). Subclass PULMONATA: Order STYLOMMATOPHORA Suborder SIGMURETHRA Infraorder ACHATINIDA Superfamily PUNCTOIDEA Family CHAROPIDAE Reticulapex Emberton & Pearce, gen. nov. Trachycystis Pilsbry, 1893 (in part), Fischer-Piette et al., 1994:175-177, 178-184, figs. 71, 73-76. Pilula Martens, 1898, Emberton, 1994a:168—171, figs. 73- 83 (not Fischer-Piette et al., 1994:193—194, figs. 85, 86). Charopidae n. gen., Emberton, 1996:730, 731, 734, 736. Type species: R. villosus sp. nov. Other species: R. apexfortis sp. nov., R. choutardi (Fi- scher-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. compactus sp. nov., R. druggi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. fischerpiettei sp. nov., R. flammulatus sp. nov., R. harananae (Emberton, 1994a) comb. nov., R. intridi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. lucidus sp. nov., R. scaber sp. nov., R. subangulatus sp. nov., R. talatai (Emberton, 1994a) comb. nov., R. ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., R. vatuvavyae (Emberton, 1994a) comb. nov., and R. vineti (Fischer- Piette, Blanc, Blanc & Salvat, 1994) comb. nov. Description: Shell diameter ca. 3-18 mm; height/diam- eter 0.6—0.8. Spire angle 100—150 degrees. Whorl periph- ery round to slightly angular. Whorl shoulder round to very flat. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome inser- tions) ca. 25—45% of shell diameter. Peristome simple, unreflected, roundish: aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.8-1.0. Distance between columellar and upper peri- stome insertions is 50-75% of aperture width. Penulti- mate whorl projecting into body whorl; occupying 4—16% of aperture height measure. Umbilicus 21-34% of shell The Veliger, Vol. 43, No. 3 diameter. Whorls of adults ca. 3.5—5.4. Coiling tightness (whorl number/In diameter) 1.9—3.3. Embryonic whorls 1.4—1.7; diameter 1.0—1.6 mm. First-whorl diameter 0.8— 1.2 mm. Embryonic sculpture obliquely crosshatched. Post-embryonic shell sculpture of transverse ribs, 7-26 ribs in the first one-tenth of the body whorl; rib height 0.5—-1.0% of shell diameter; rib periostracum without or- namentation or locally extended into hairs. Peripheral, non-rib periostracum without ornamentation or locally extended into hairs. Spiral sculpture absent or restricted to a few spiral grooves below the periphery of the body whorl. Shell basic color yellow-brown to brown-red. Sec- ondary coloration absent or consisting of whitish flam- mulations or blotches. Penis length 0.4—0.9 shell diameter, thick, slightly bul- bous apically. Penis without sheath or caecum. Penial sculpture consisting of various combinations of smooth (rarely pustulose), soft bulges, flaps, pilasters, and ridges. Epiphallus a bulbous sac arising subterminally from the apical penial bulb; epiphallar bulb lying alongside, and tightly adherent to, the apical penial bulb. Spermatophore less than three times as long as wide, tapered, spiraling slightly; spermatophore attaches to the mate’s basal penis. Penial retractor muscle attached to the epiphallar bulb, below the bulb’s apex. Vas deferens enters the epiphallus basally, near the epiphallus’s juncture with the penis; vas deferens slender along its entire length. Atrium short to long. Spermathecal duct joining the oviduct near the ovi- duct’s entry to the atrium, thus the vagina is short. Sper- mathecal duct slender; spermatheca unknown. Fertiliza- tion pouch-seminal receptacle complex extremely long and slender, generally free of the albumen gland. Diagnosis: Reticulapex gen. nov. seems to be a mono- phyletic clade diagnosed by, and unique for, its obliquely cross-hatched embryonic sculpture; an extremely elon- gate fertilization pouch-seminal receptable complex; nearly basal entry of the vas deferens into, and nearly apical origin of the penial-retractor muscle from, the epi- phallus; and a spiral spermatophore that is transferred to the mate’s basal penis. Comments: Reticulapex gen. nov. and Trachycystis Pils- bry, 1893, seem to be the only genera of Charopidae known so far in Madagascar. We have collected exten- sively in Tsaratanana Reserve and found species very similar to ‘‘Pilula’’ excavata Fischer-Piette, Blanc & Sal- vat, 1975, and ‘‘Pilula’’ madecassina Fischer-Piette, Blanc & Salvat, 1975, of the same massif; they are not Pilula Martens, 1898, but seem to be Helicarionidae, ten- tatively Microcystis Beck, 1837 (Emberton, unpublished). Surprisingly, we did not find any Charopidae in Tsaratan- ana Reserve (Emberton, unpublished). ‘“*Pilula’”’ harananae Emberton, 1994; ‘‘Pilula”’ talatai Emberton, 1994; and ‘‘Pilula’’ vatuvavyae Emberton, 1994, are not Pilula, but Reticulapex gen. nov. Trachycystis is currently assigned to Charopidae K. C. Emberton & T. A. Pearce, 2000 (Vaught, 1989), not Punctidae as erroneously stated by Fischer-Piette et al. (1994). Only one of Fischer-Piette et al.’s (1994) ‘“‘Trachycystis’> seems validly placed: Tra- chycystis waterloti Fischer-Piette, Blanc, Blanc & Salvat, 1994. Their other species of ‘“‘Trachycystis’’ are either Reticulapex gen. nov. (five species listed above) or, in all likelihood, Helicarionidae, tentatively either Microcystis or Sitala Adams, 1865 (Emberton, unpublished). The spermatophore is known so far only in R. com- pactus sp. nov. and in the type species R. villosus sp. nov. Reticulapex subangulatus Emberton & Pearce, sp. nov. (Figures 1, 13, 18) Charopidae sp. 1, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860810 (ex MBI 383.02DH, Tol-11, ad). Paratypes: MBI 380.14DP (4 ad, 6 juv), MBI 382.15DP (1 ad, 5 juv; AMS C.203454 [1 ad]; MNHN [1 ad]; ANSP 400836 [1 ad], MBI 382.15AP (2 ad [1 dissected], 1 juv), MBI 383.02DP (2 ad, 4 juv), MBI 384.14DP (2 ad, 3 juv), MBI 384.14AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: southeast slope of Mount Vasiha, 700 m elevation: latitude 24°55'23"S, longitude 46°44'27’E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 7.8 mm; height 4.6 mm. Height-diameter ratio 0.6. Spire angle 130 degrees. Whorl periphery slightly angular. Whorl shoulder very flat. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 36% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 1.0. Distance between columellar and upper peristome in- sertions is 61% of aperture width. Penultimate whorl pro- jecting into body whorl; occupying 11% of aperture height measure. Umbilicus 26% of shell diameter. Whorls 4.1. Coiling tightness 2.0. Embryonic Shell. Whorls 1.4; diameter 1.2 mm. First- whorl diameter 1.0 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 7.7. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture very faint. Transverse rib density 10 in first tenth of body whorl. Rib height 1.0% of shell diameter. Rib per- iostracum without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture absent or incon- spicuous. Shell basic color yellow-brown. Secondary col- oration absent. Page 251 Shell variation: Occasional shells have a slightly less angular periphery. Shell comparisons: Most similar to Reticulapex druggi comb. nov., but more than 50% larger with fewer whorls. Description of lower reproductive system (MBI 382.15AP: 1 adult): Penis 4.7 mm long (0.6 shell di- ameter), 1.0—1.4 mm wide, slightly bulbous apically. Pe- nis without sheath or caecum. Penial sculpture consisting of a long apical flap (ca. 2.0 mm unfurled) and an elon- gate lateral bulge (ca. 1.7 mm long). Epiphallus a bulbous sac arising subterminally from the apical penial bulb; epi- phallar bulb lying alongside, and tightly adherent to, the apical penial bulb. Penial retractor muscle attached to the epiphallar bulb, below the bulb’s apex. Vas deferens en- ters the epiphallus basally, near the epiphallus’s juncture with the penis; vas deferens slender along its entire length. Atrium short. Spermathecal duct joining the ovi- duct near the oviduct’s entry to the atrium, thus the va- gina is short. Spermathecal duct slender and long; sper- matheca ovoid, lying alongside the proximal albumen gland. Fertilization pouch-seminal receptacle complex ex- tremely long (ca. 2.5 mm) and slender, free of the albu- men gland. Distribution and conservation status: Mts. [lapiry and Vasiha, 300 to 860 m elevation. Not known from any other localities (Emberton, unpublished). Thus, occurring as fragmented subpopulations within < 1000 km? of de- clining forest in the southern Anosy and Vohimena Chains, therefore, by IUCN (1996) criteria, an Endan- gered species. Etymology: For the sub-angulate periphery. Reticulapex lucidus Emberton & Pearce, sp. nov. (Figures 2, 14, 19) Charopidae n. gen. u/richi Fischer-Piette et al., 1994, Em- berton, 1996:736 (due to a printer’s error replacing “‘n. sp. 13,” rather than the correct “n. sp. 14,” with ‘‘ul- richi’’). Charopidae sp. 2, Emberton et al., 1996:210. Emberton, 1997:1146, 1149. Emberton et al., 1999:table 2. Holotype: USNM 860811 (ex MBI 378.03DH, Tol-6, ad). Paratypes: MBI 377.15DP (10 ad, 30 juv), MBI 377. 15AP (13 juv), MBI 378.03DP (10 ad, 21 juv; AMS C.203455 [1 ad]; MNHN [1 ad]; ANSP 400837 [1 ad]), MBI 378.03AP (1 ad [dissected], 13 juv), MBI 379.22DP (4 ad, 9 juv), MBI 379.22AP (6 juv), MBI 380.15DP (6 ad, 10 juv), MBI 380.15AP (3 ad, 5 juv), MBI 381.15DP (8 ad, 17 juv), MBI 381.15AP (2 ad, 4 juv), MBI 382.16DP (2 juv), MBI 382.16AP (1 juv), MBI 388.09DP (1 juv), MBI 391.06AP (1 ad). Type locality: Madagascar: Tulear Province: northwest 52 The Veliger, Vol. 43, No. 3 Figures 1-3 Figure |. Reticulapex subangulatus Emberton & Pearce, sp. nov., holotype. Figure 2. Reticulapex lucidus Emberton & Pearce, sp. nov., holotype. Figure 3. Reticulapex scaber Emberton & Pearce, sp. nov., holotype. All scale bars | mm. K. C. Emberton & T. A. Pearce, 2000 Page 253 of Fort Dauphin: west of village of Mahialambo: ridge on east slope of Mt. Ilapiry, 500 m elevation, latitude 24°51'33”S, longitude 47°00'27’E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 6.6 mm; height 3.9 mm. Height-diameter ratio 0.6. Spire angle 140 degrees. Whorl periphery round. Whorl shoulder round. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 41% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.9. Dis- tance between columellar and upper peristome insertions is 63% of aperture width. Penultimate whorl projecting into body whorl; occupying 13% of aperture height mea- sure. Umbilicus 25% of shell diameter. Whorls 4.3. Coil- ing tightness 2.3. Embryonic Shell. Whorls 1.7; diameter 1.2 mm. First- whorl diameter 1.1 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 9.3. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture very faint. Transverse rib density 15 in first tenth of body whorl. Rib height 0.5% of shell diameter. Rib per- iostracum without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture conspicuous; 10 grooves below periphery of body whorl. Shell basic color brown-yellow. Secondary coloration absent. Shell variation: Some shells have a noticeably higher spire than the holotype. Shell comparisons: Much tighter coiling than Reticula- pex ulrichi comb. nov. Description of lower reproductive system (MBI 378.03AP: 1 adult): Penis 3.8 mm long (0.6 shell di- ameter), 0.8—1.1 mm wide, slightly bulbous apically. Pe- nis without sheath or caecum. Penial sculpture consisting of a long apical flap (ca. 1.7 mm unfurled); an elongate lateral bulge (ca. 1.5 mm long); and a parallel, crestlike pilaster (ca. 2.0 mm long). Epiphallus a bulbous sac aris- ing subterminally from the apical penial bulb; epiphallar bulb lying alongside, and tightly adherent to, the apical penial bulb. Penial retractor muscle attached to the epi- phallar bulb, below the bulb’s apex. Vas deferens enters the epiphallus basally, near the epiphallus’s juncture with the penis; vas deferens slender along its entire length. Atrium long. Spermathecal duct joining the oviduct near the oviduct’s entry to the atrium, thus the vagina is short. Spermathecal duct slender and long; spermatheca ovoid, lying alongside the proximal albumen gland. Fertilization pouch-seminal receptacle complex extremely long (ca. 2.2 mm) and slender, free of the albumen gland. Distribution and conservation status: Mts. Ilapiry and Vasiha, 200 to 860 m elevation. Also reported from An- dohahela (1200-1960 m) in the Anosy Chain, and from Mt. Teloboko (640 m, just below the summit), which is near Mt. Mahermana in the Vohimena Chain; but from nowhere else (Emberton, unpublished). Thus an Endan- gered species by IUCN (1996) criteria: restricted to < 2000 km? of forest that is continually declining in extent and quality, and occurring as isolated subpopulations. Etymology: For the shell surface being relatively shiny (L. lucid-) for the genus. Reticulapex scaber Emberton & Pearce, sp. nov. (Figure 3) Charopidae n. gen. n. sp. 12, Emberton, 1996:736. Charopidae sp. 3, Emberton et al., 1996:210. Emberton 1997:1147, 1150. Emberton et al., 1999:table 2. Holotype: USNM 860812 (ex MBI 376.04DH, Tol-4, ad). Paratypes: MBI 374.16DP (2 juv), MBI 375.14DP (1 ad, 2 juv; AMS C.203456 [1 ad]), MBI 376.04DP (2 juv), MBI 376.04AP (1 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: valley on northwest slope of Mt. Mahermana, 100 m elevation: lat- itude 24°26'22"S, longitude 47°12'41”E: primary rainfo- rest. Description of holotype shell: Shell Size and Shape. Diameter 8.6 mm; height 6.6 mm. Height-diameter ratio 0.8. Spire angle 110 degrees. Whorl periphery round. Whorl shoulder flat. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 41% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 1.0. Dis- tance between columellar and upper peristome insertions is 49% of aperture width. Penultimate whorl projecting into body whorl; occupying 9% of aperture height mea- sure. Umbilicus 23% of shell diameter. Whorls 4.9. Coil- ing tightness 2.3. Embryonic Shell. Whorls 1.6; diameter 1.4 mm. First- whorl diameter 0.9 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 4.8. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture strong. Transverse rib density 7 in first tenth of body whorl. Rib height 1.6% of shell diameter. Rib periostrac- um without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture absent or inconspicuous. Shell basic color brown-yellow. Secondary coloration ab- sent. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Most similar to Reticulapex vineti comb. nov., but larger for the same number of whorls and lacking a caniculate suture and flamulate coloration. Distribution and conservation status: Mt. Mahermana, 100 to 300 m elevation. Also reported from the nearby Mts. Teloboko (640 m) and Esetra (mid-elevational slope), and from the central-Vohimena-chain Mt. Varabe (200 m), but nowhere else (Emberton, unpublished). Thus this spe- cies appears to be restricted to the northern Vohimena Mountain Chain. Its forest habitat covers < 400 km* that is diminishing rapidly in area and quality, and it is frag- mented into isolated subpopulations with reduced proba- bility of recolonization. By IUCN (1996) criteria, Reticu- lapex gen. nov. scaber sp. nov. is Endangered. Etymology: For the rough (L. scaber) surface sculpture. Reticulapex compactus Emberton & Pearce, sp. nov. (Figures 4, 15, 20) Charopidae sp. 4, Emberton et al., 1996:210. Emberton, 1997:1146, 1150. Emberton et al., 1999:table 2. Holotype: USNM 860813 (ex MBI 373.08DH, Tol-1, ad). Paratypes: MBI 373.08DP (1 ad, 1 juv), MBI 373.08AP (2 ad [1 dissected], 3 juv), MBI 374.17DP (2 ad, 7 juv; AMS C.203457 [1 ad]; MNHN [1 ad]; ANSP 400838 [1 ad]), MBI 375.15DP (5 ad, 3 juv), MBI 375.15AP (1 ad, 2 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: summit of Mt. Mahermana, 340 m elevation: latitude 24°26'12”S, longitude 47°13'13”E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 3.3 mm; height 2.6 mm. Height-diameter ratio 0.8. Spire angle 100 degrees. Whorl periphery round. Whorl shoulder round. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 42% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.8. Dis- tance between columellar and upper peristome insertions is 75% of aperture width. Penultimate whorl projecting into body whorl; occupying 10% of aperture height mea- sure. Umbilicus 23% of shell diameter. Whorls 3.9. Coil- ing tightness 3.3. Embryonic Shell. Whorls 1.5; diameter 1.0 mm. First- whorl diameter 0.8 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 15.7. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture very faint. Transverse rib density 9 in first tenth of The Veliger, Vol. 43, No. 3 body whorl. Rib height 0.6% of shell diameter. Rib per- iostracum without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture absent or incon- spicuous. Shell basic color brown. Secondary coloration absent. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Most similar to Reticulapex kerma- deci comb. nov., but twice as large for the same number of whorls and with a proportionally smaller aperture. Description of lower reproductive system (MBI 373.08AP: 1 adult): Penis 2.0 mm long (0.6 shell di- ameter), 0.4—0.6 mm wide, slightly bulbous apically. Pe- nis without sheath or caecum. Penial sculpture consisting of a thin apical pilaster (ca. 0.7 mm long), below which is an ovoid lateral bulge (ca. 0.7 mm long). Epiphallus a bulbous sac arising subterminally from the apical penial bulb; epiphallar bulb lying alongside, and tightly adherent to, the apical penial bulb. Spermatophore consisting of an egg-shaped body (ca. 0.6 mm long) with a stalklike pro- cess (ca. 0.6 mm long) that spirals slightly; spermato- phore found attached within the basal penial tube. Penial retractor muscle attached to the epiphallar bulb, below the bulb’s apex. Vas deferens enters the epiphallus basal- ly, near the epiphallus’s juncture with the penis; vas de- ferens slender along its entire length. Atrium long. Sper- mathecal duct joining the oviduct near the oviduct’s entry to the atrium, thus the vagina is short. Spermathecal duct slender and long; spermatheca ovoid, lying alongside the proximal albumen gland. Fertilization pouch-seminal re- ceptacle complex unknown. Distribution and conservation status: Mt. Mahermana, 200 to 340 m elevation. Also reported from a nearby patch of coastal forest at about 15 m elevation. This is a Critically Endangered species, by criteria of the IUCN (1996), because it is highly fragmented between two patches within < 100 km? of forest habitat that is contin- ually declining in area and quality. Etymology: For the compact shape. Reticulapex flammulatus Emberton & Pearce, sp. nov. (Figure 5) Charopidae sp. 5, Emberton et al., 1996:210. Emberton, 1997:1146, 1150. Emberton et al., 1999:table 2. Holotype: USNM 860814 (ex MBI 377.03DH, Tol-5, ad). Paratypes: MBI 373.17DP (1 juv), MBI 374.18DP (1 juv), MBI 375.16DP (3 juv), MBI 377.03DP (1 juv), MBI 377.03AP (4 juv), MBI 378.15DP (1 ad, 1 juv; AMS C.203458 [1 ad]; MNHN [1 ad]), MBI 379.23DP (1 ad). K. C. Emberton & T. A. Pearce, 2000 Page 255 Figures 4 and 5 Figure 4. Reticulapex compactus Emberton & Pearce, sp. nov., holotype. Figure 5. Reticulapex flammulatus Em- berton & Pearce, sp. nov., holotype. All scale bars 1 mm. Page 256 The Veliger, Vol. 43, No. 3 Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Mahialambo: summit of Mt. Ilapiry, 540 m elevation: latitude 24°51'40"S, lon- gitude 47°00'20"E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 6.5 mm; height 3.6 mm. Height-diameter ratio 0.6. Spire angle 150 degrees. Whorl periphery round. Whorl shoulder round. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 40% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.9. Dis- tance between columellar and upper peristome insertions is 55% of aperture width. Penultimate whorl projecting into body whorl; occupying 10% of aperture height mea- sure. Umbilicus 34% of shell diameter. Whorls 3.8. Coil- ing tightness 2.0. Embryonic Shell. Whorls 1.6; diameter 1.6 mm. First- whorl diameter 1.1 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 3.4. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture faint. Transverse rib density 14 in first tenth of body whorl. Rib height 0.6% of shell diameter. Rib periostrac- um without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture absent or inconspicuous. Shell basic color brown-red. Secondary coloration white blotches. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Larger, flatter, and more loosely coiled than Reticulapex intridi comb. nov. Fewer whorls (but same size) and much stronger ribbed sculpture than Reticulapex vatuvavyae comb. nov. Distribution and conservation status: Mts. Mahermana and [apiry, 200 to 540 m elevation. Not known from any other localities, not even from several peaks adjacent to Mt. Mahermana (Emberton, unpublished). Clearly an En- dangered species (IUCN [1996] criteria), restricted to fragmented subpopulations within the Vohimena Chain, whose forests extend only about 500 km? and are contin- ually declining. Etymology: For the flame-colored (L. flamma) markings on the shell. Reticulapex fischerpiettei Emberton & Pearce, sp. nov. (Figure 6) Charopidae n. gen. n. sp. 10, Emberton, 1996:736. Charopidae sp. 6, Emberton et al., 1996:210. Emberton, 1997:1146, 1150. Emberton et al., 1999:table 2. Holotype: USNM 860815 (ex MBI 379.03DH, Tol-7, ad). Paratypes: MBI 379.03DP (2 ad, 2 juv; AMS C.203459 [1 ad]; MNHN [1 ad]; ANSP 400839 [1 ad]), MBI 379.03AP (2 juv), MBI 380.16DP (1 ad, 4 juv), MBI 381.16DP (3 ad, 7 juv), MBI 381.16AP (3 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Mahialambo: ridge, val- ley, and slope on southsoutheast slope of Mt. Hapiry, 400m elevation: latitude 24°51'27’S, longitude 47°00'38’E: pri- mary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 3.9 mm; height 2.8 mm. Height-diameter ratio 0.7. Spire angle 100 degrees. Whorl periphery slightly angular. Whorl shoulder very flat. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 38% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.9. Distance between columellar and upper peristome in- sertions is 54% of aperture width. Penultimate whorl pro- jecting into body whorl; occupying 4% of aperture height measure. Umbilicus 23% of shell diameter. Whorls 3.8. Coiling tightness 2.8. Embryonic Shell. Whorls 1.5; diameter 1.2 mm. First- whorl diameter 0.9 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 8.2. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture moderate. Transverse rib density 8 in first tenth of body whorl. Rib height 1.0% of shell diameter. Rib per- iostracum locally extended into hairs; hair length 12.8% of shell diameter. Non-rib periostracum without ornamen- tation. Spiral sculpture absent or inconspicuous. Shell ba- sic color brown-red. Secondary coloration absent. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Unique within the genus for its len- ticular shape and long peripheral hairs. Distribution and conservation status: Mt. Ilapiry, 200 to 400 m elevation. Also reported (Emberton, unpub- lished) from the nearby Mt. St. Jacques (520 m) and, in the Anosy Chain, from Andohahela (1400-1600 m); but nowhere else. Thus fragmented within the southern Voh- imena and Anosy forests, whose extent is < 1000 km? and is continually declining; therefore, by IUCN (1996) standards, an Endangered species. Etymology: For the late E. Fischer-Piette, longtime sys- tematist of Madagascar’s land snails. K. C. Emberton & T. A. Pearce, 2000 Page i Figures 6-8 Figure 6. Reticulapex fischerpiettei Emberton & Pearce, sp. nov., holotype. Figure 7. Reticulapex apexfortis Em- berton & Pearce, sp. nov., holotype. Figure 8. Reticulapex ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., representative from Mt. Vasiha at 100 m elevation. All scale bars | mm. Page 258 Reticulapex apexfortis Emberton & Pearce, sp. nov. (Figure 7) Charopidae sp. 7, Emberton et al., 1996:210. Emberton, 1997:1147. Emberton et al., 1999:table 2. Holotype: USNM 860816 (ex MBI 384.03DH, Tol-12, ad). Paratypes: MBI 384.03DP (1 juv), MBI 384.03AP (2 juv). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: west of village of Malio: east slope of Mt. Vasiha, 500 m elevation: latitude 24°55'19"S, longi- tude 46°44'45”E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 3.6 mm; height 2.2 mm. Height-diameter ratio 0.6. Spire angle 140 degrees. Whorl periphery round. Whorl shoulder round. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 43% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.9. Dis- tance between columellar and upper peristome insertions is 49% of aperture width. Penultimate whorl projecting into body whorl; occupying 16% of aperture height mea- sure. Umbilicus 26% of shell diameter. Whorls 3.2. Coil- ing tightness 2.5. Embryonic Shell. Whorls 1.5; diameter 1.3 mm. First- whorl diameter 0.8 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 6.5. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture faint. Transverse rib density 9 in first tenth of body whorl. Rib height 0.8% of shell diameter. Rib periostrac- um without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture absent or inconspicuous. Shell basic color brown. Secondary coloration absent. Shell comparisons: Similar to Reticulapex choutardi comb. nov., but without its complex embryonic sculpture, without its spiral sculpture, and with a higher spire (height/diameter 0.8 as opposed to 0.6). Distribution and conservation status: Known only from Mt. Vasiha at 500 m elevation. No records elsewhere (Emberton, unpublished). By IUCN (1996) criteria (sin- gle locality, < 100 km? extent, continuing decline in area and quality of habitat), Reticulapex gen. nov. apexfortis sp. nov. is Critically Endangered. Etymology: For the strong (L. fortis) sculpturing on its embryonic whorls, or tip (L. apex). The Veliger, Vol. 43, No. 3 Reticulapex ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov. (Figures 8, 16, 21) Trachycystis ulrichi n. sp., Fischer-Piette et al., 1994:175— ipetiondAl Charopidae n. gen. n. sp. 3, Emberton, 1996:736. Charopidae n. gen. n. sp. 14 (not “‘n. gen. ulrichi,”> which was a printer’s error), Emberton, 1996:736. Charopidae sp. 8, Emberton et al., 1996:210. Emberton, 1997:1147. Emberton et al., 1999:table 2. Representative: MBI 388.01DR, Tol-16 (ad). Other specimens: MBI 388.01A (1 ad [dissected]). Description of representative: Shell Size and Shape. Diameter 5.4 mm; height 3.3 mm. Height-diameter ratio 0.6. Spire angle 140 degrees. Whorl periphery round. Whorl shoulder round. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 46% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 0.9. Dis- tance between columellar and upper peristome insertions is 50% of aperture width. Penultimate whorl projecting into body whorl; occupying 12% of aperture height mea- sure. Umbilicus 25% of shell diameter. Whorls 3.3. Coil- ing tightness 2.0. Embryonic Shell. Whorls 1.5; diameter 1.5 mm. First- whorl diameter 1.1 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 3.7. Embryonic sculpture crosshatched. Shell Sculpture and Color. Neanic cross-hatch sculp- ture absent (undetected). Transverse rib density 8 in first tenth of body whorl. Rib height 0.2% of shell diameter. Rib periostracum without ornamentation. Non-rib perios- tracum without ornamentation. Non-rib periostracum without ornamentation. Spiral sculpture absent or incon- spicuous. Shell basic color brown-yellow. Secondary col- oration absent. Description of lower reproductive system (MBI 388.01A: 1 adult): Penis 5.0 mm long (0.9 shell diam- eter), 0.8—1.2 mm wide, slightly bulbous apically. Penis without sheath or caecum. Penial sculpture consisting of two parallel apical pilasters (ca. 2.5 mm long), below one of which is an ovoid lateral bulge (ca. 1.5 mm long). Epiphallus a bulbous sac arising subterminally from the apical penial bulb; epiphallar bulb lying alongside, and tightly adherent to, the apical penial bulb. Penial retractor muscle attached to the epiphallar bulb, below the bulb’s apex. Vas deferens enters the epiphallus basally, near the epiphallus’s juncture with the penis; vas deferens slender along its entire length. Atrium short. Spermathecal duct joining the oviduct near the oviduct’s entry to the atrium, K. C. Emberton & T. A. Pearce, 2000 thus the vagina is short. Spermathecal duct slender and long; spermatheca ovoid, lying alongside the proximal albumen gland. Fertilization pouch-seminal receptacle complex extremely long (ca. 5 mm) and slender. Distribution and conservation status: Mt. Vasiha at 100 m elevation. Described (Fischer-Piette et al., 1994) from elsewhere in the Anosy Chain, 1000-1950 m. Emberton (unpublished) gives records from Andohahela (450-1900 m), Col Beampingaratra (380-1280 m), Mt. Ramabeafo (410-700 m), and other localities in the southern Anosy Chain and in the ridge connecting it to the southern Voh- imena Chain; as well as in the northern, but not the south- ern, Vohimena Chain: Mt. Mahermana (300 m) and Mt. Varabe (200 m). With a range restricted to the Anosy and northern Vohimena Chains, this species is Endangered, under IUCN criteria (IUCN, 1996: occurrence < 5000 km’, severely fragmented, habitat extent and quality in continuing decline). Reticulapex villosus Emberton & Pearce, sp. nov. Crimes DO, il, WA, 15 22, 28) Charopidae n. gen. n. sp. 11, Emberton, 1996:736. Charopidae sp. 9, Emberton et al., 1996:210. Emberton, 1997:1147, 1150. Emberton et al., 1999:table 2. Holotype: USNM 860817 (ex MBI 389.01DH, Tol-3-4, ad). Paratypes: MBI 373.26AP (1 ad), MBI 374.25AP (2 juv), MBI 375.22AP (3 ad [1 dissected], 2 juv), MBI 389.01AP (2 ad [1 dissected]), MBI 390.02DP (0; AMS C.203460 [1 ad]). Type locality: Madagascar: Tulear Province: northwest of Fort Dauphin: northeast of village of Esetra: west slope of Mt. Mahermana, 100—200 m elevation, latitude 24°26'15-22”S, longitude 47°13'04—-12".41E: primary rainforest. Description of holotype shell: Shell Size and Shape. Diameter 18.1 mm; height 10.2 mm. Height-diameter ratio 0.6. Spire angle 140 degrees. Whorl periphery round. Whorl shoulder flat. Aperture width (inside dimension, parallel to a line between the columellar and upper peristome insertions) 38% of shell diameter. Aperture height-width ratio (inside dimension, height measured to and perpendicular to a line between the columellar and upper peristome insertions) 1.0. Dis- tance between columellar and upper peristome insertions is 64% of aperture width. Penultimate whorl projecting into body whorl; occupying 13% of aperture height mea- sure. Umbilicus 21% of shell diameter. Whorls 5.4. Coil- ing tightness 1.9. Embryonic Shell. Whorls 1.7; diameter 1.6 mm. First- whorl diameter 1.2 mm. Coiling tightness (embryonic whorl number divided by natural logarithm of embryonic shell diameter) 3.6. Embryonic sculpture crosshatched. Page 259 Shell Sculpture and Color. Neanic cross-hatch sculp- ture absent (undetected). Transverse rib density 26 in first tenth of body whorl. Rib height 0.1% of shell diameter. Rib periostracum without ornamentation. Non-rib perios- tracum with bristles; 18 bristles in each inter-rib space between sutures at end of penultimate whorl; bristle length 1.3% of shell diameter. Spiral sculpture absent or inconspicuous. Shell basic color brown-red. Secondary coloration absent. Shell variation: No conspicuous variation in size or shape. Shell comparisons: Unique within the genus for its huge size and dense hairs. Description of lower reproductive system (MBI 375.22AP: 1 adult; MBI 389.01AP: 1 adult): Penis 7.8 mm long (0.4 shell diameter), 1.4—2.5 mm wide, slightly bulbous apically. Penis without sheath or caecum. Penial sculpture consisting of an apical, ovoid, pustulose (glan- dular?) bulb (ca. 1.7 mm long); a thick, V-shaped pilaster in the upper half of the penis (length ca. 3.9 mm); a field of basal ridges that are low, parallel, and even (ca. 14 in number); and an indistinct basal pilaster (ca. 4.7 mm long). Epiphallus a bulbous sac arising subterminally from the apical penial bulb; epiphallar bulb lying along- side, and tightly adherent to, the apical penial bulb. Sper- matophore thick-bodied, tapered apically, spiraling slight- ly (ength ca. 3.4 mm); spermatophore found attached within the basal penial tube. Penial retractor muscle at- tached to the epiphallar bulb, below the bulb’s apex. Vas deferens enters the epiphallus basally, near the epiphal- lus’s juncture with the penis; vas deferens slender along its entire length. Atrium short. Spermathecal duct joining the oviduct near the oviduct’s entry to the atrium, thus the vagina is short. Spermathecal duct slender; sperma- theca unknown. Fertilization pouch-seminal receptacle complex extremely long (total length unknown) and slen- der, free of the albumen gland. Distribution and conservation status: Mt. Mahermana, 200 to 340 m elevation. Also found on Mt. Varabe at 410 m, but nowhere else (Emberton, unpublished). Apparently restricted to the northern Vohimena Mountain Chain, fragmented into subpopulations within dwindling forest < 500 km? in extent. Therefore, by IUCN (1996) criteria, an Endangered species. Etymology: For the hairy (L. villos-) shell surface. CHAROPID CONSERVATION STATUSES Analyses of individual species are given above in the spe- cies descriptions. To summarize, all nine of the charopid species are proposed as either Endangered or Critically Endangered. Two species should be listed as Critically Endangered: R. apexfortis sp. nov. and Reticulapex compactus sp. nov. The seven species that should be list- Page 260 The Veliger, Vol. 43, No. 3 WA) iC 1 Figures 9-11 Reticulapex villosus Emberton & Pearce, sp. nov. Figure 9. Holotype. Figure 10. Paratype, MBI 390.02DP. Figure 11. Other paratype, MBI 375.22AP. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 Page 261 or oe Figure 12 Some anatomical characters used in descriptions, as shown on Reticulapex villosus Emberton & Pearce, sp. nov. Abbreviations: A-albumen gland, B-pustulate bulb (penial sculpture), D-hermaphroditic duct, E-epiphallus, F-fer- tilization pouch-seminal receptacle complex (broken short in this specimen), G-genital pore, M-penial retractor muscle, P-penis, R-parallel ridges (penial sculpture), T-spermatophore (with deposition site), U-prostate-uterus, V- vas deferens. ed as Endangered are: R. fischerpiettei sp. nov., R. flam- mulatus sp. nov., R. lucidus sp. nov., R. scaber sp. nov., R. subangulatus sp. nov., R. ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov., and R. villosus sp. nov. DISCUSSION These descriptions of nine charopids support our prior distributional and ecological analyses of Mahermana-ll- apiry-Vasiha land snails (Emberton et al., 1996, 1999; Emberton, 1997). Two previous papers (Emberton & Pearce, 1999, 2000) dealt with the caenogastropods and the small, high-spired pulmonates. A fourth and final pa- per in this series is in press describing the Mahermana- Ilapiry-Vasiha helicarionids. The Madagascan charopid radiation—contrary to an early prediction (Emberton, 1994a)—-seems to be concen- trated in the southeastern rainforests, with diversity drop- ping off steeply toward the north and with a total of 24 known species (Emberton, unpublished). Further explo- ration, particularly in the southeast, should yield many more species. Such exploration must be prompt, as much of the most diverse rainforest of the region will be soon be eradicated (Emberton, 1997; Emberton et al., 1999). Page 262 The Veliger, Vol. 43, No. 3 Figures 13-16 Charopid reproductive systems. Figure 13. Reticulapex subangulatus Emberton & Pearce, sp. nov. Figure 14. Reticulapex lucidus Emberton & Pearce, sp. nov. Figure 15. Reticulapex compactus Emberton & Pearce, sp. nov. Figure 16. Reticulapex ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov. All scale bars 1 mm. K. C. Emberton & T. A. Pearce, 2000 Page 263 Figures 17—23 Charopid reproductive systems, penes, and spermatophores. Figure 17. Reticulapex villosus Emberton & Pearce, sp. nov., MBI 375.22AP (right and left) and MBI 389.01AP (middle). Figure 18. Reticulapex subangulatus Em- berton & Pearce, sp. nov. Figure 19. Reticulapex lucidus Emberton & Pearce, sp. nov. Figure 20. Reticulapex compactus Emberton & Pearce, sp. nov. Figure 21. Reticulapex ulrichi (Fischer-Piette, Blanc, Blanc & Salvat, 1994) comb. nov. Figures 22, 23, Reticulapex villosus Emberton & Pearce, sp. nov., MBI 375.22AP (Figure 22) and MBI 389.01AP (Figure 23). All scale bars 1 mm. Page 264 The Veliger, Vol. 43, No. 3 It seems very likely that that all but one (Trachycystis waterloti) of Madagascar’s charopids belong to the en- demic genus Reticulapex gen. nov. Fischer-Piette et al.’s (1994) ‘‘Pilula’’? seems to be helicarionid and should probably be transferred to Microcystis (this paper; Em- berton, unpublished). Reticulapex gen. nov. seems to be a monophyletic clade defined by cross-hatched embry- onic sculpture, a long FPSC, a distinctive epiphallus, and a spiral spermatophore that is transferred to the mate’s basal penis. Spermatophore placement on the mate’s penis is known to occur only by external sperm exchange. This is a de- rived condition in which two mating snails evert and en- twine, rather than insert, penes. External sperm exchange is apparently unique to stylommatophorans, within which it evolved independently in the Limacidae, Discidae, Suc- cineidae, and Polygyridae (review in Emberton, 1994b). Thus Reticulapex gen. nov. represents the fifth known independent evolution of external sperm exchange within the Stylommatophora. It is clear that external sperm ex- change is not a general phenomenon within the Charo- pidae. Dissected charopids from Australia (Stanisic, 1990, 1993) and New Caledonia (Mordan & Tillier, 1986) have spermatophores within the female tract, implying internal sperm exchange. Acknowledgments. We are grateful to the U.S. National Science Foundation and USAID for funding (grant DEB-9201060 to KCE); to staffs of the Ranomafana National Park Project, the Madagascar Départment des Eaux et Foréts, and the Tolagnaro (Fort Dauphin) office of the World Wide Fund for Nature for logistical aid; to Roger Randalana and assistants from Esetra, Mahialambo, and Malio for collecting; to the late Felix Rako- tomalala for curatorial assistance; and to Lucia Emberton for help in mounting the photographs. LITERATURE CITED Da.tLwitz, M. J., T. A. PAINE & E. J. ZURCHER. 1993. DELTA User’s Guide: A General System for Processing Taxonomic Descriptions. 4th ed. CSIRO Information Services: Mel- bourne, Australia. 136 pp. EMBERTON, K. C. 1994a. Thirty new species of Madagascan land snails. Proceedings of the Academy of Natural Sciences of Philadelphia 145:147—189. EMBERTON, K. C. 1994b. Polygyrid land-snail phylogeny: exter- nal sperm exchange, early North American biogeography, iterative shell evolution. Biological Journal of the Linnean Society 52:241—271. EMBERTON, K. C. 1996. Conservation priorities for forest-floor invertebrates of the southeastern half of Madagascar: evi- dence from two land-snail clades. Biodiversity and Conser- vation 5:729-741. EMBERTON, K. C. 1997. Diversities, distributions, and abundances of 80 species of minute-sized land snails in southeastern- most Madagascan rainforests, with a report that lowlands are richer than highlands in endemic and rare species. Biodi- versity and Conservation 6:1137—1154. EMBERTON, K. C. & T. A. PEARCE. 1999. Land caenogastropods from Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with conservation statuses of 17 Boucardicus. The Veliger 42:338-372. EMBERTON, K. C. & T. A. PEARCE. 2000. Small, high-spired pul- monates from Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with description of a new genus, and with conservation statuses of 15 streptaxid species. The Veliger 43:126-153. EMBERTON, K. C., T. A. PEARCE & R. RANDALANA. 1996. Quan- titatively sampling land-snail species richness in Madagas- can rainforests. Malacologia 38:203-212. EMBERTON, K. C., T. A. PEARCE & R. RANDALANA. 1999. Mol- luscan diversity in the unconserved Vohimena and the con- served Anosy mountain chains, southeast Madagascar. Bio- logical Conservation 89:183-188. FISCHER-PIETTE, E., C. P- BLANC, EF BLanc & E Satvart. 1994. Gastéropodes terrestres pulmonés. Faune de Madagascar 83: 1-551. GREEN, G. M. & R. W. SussMANn. 1990. Deforestation history of the eastern rain forests of Madagascar from satellite images. Science 248:212-215. IUCN. 1996. 1996 IUCN Red List of Threatened Animals. In- ternational Union for the Conservation of Nature, Gland, Switzerland. 368 pp. Morpan, P. & S. TILLIER. 1986. New Caledonian charopid land snails. I. Revision of the genus Pararhytida (Gastropoda: Charopidae). Malacologia 27:203—241. NorpbsIiEck, H. 1986. The system of the Stylommatophora (Gas- tropoda), with special regard to the systematic position of the Clausiliidae, II. Importance of the shell and distribution. Archiv fiir Molluskenkunde 117:93-116. PONDER, W. E & D. R. LINDBERG. 1997. Towards a phylogeny of gastropod molluscs: an analysis using morphological characters. Zoological Journal of the Linnean Society 119: 83-265. STANISIC, J. 1990. Systematics and biogeography of eastern Aus- tralian Charopidae (Mollusca, Pulmonata) from subtropical rainforests. Memoirs of the Queensland Museum 30:1—241. STANISIC, J. 1993. Danielleilona gen. nov., from the wet tropics, northeastern Queensland (Pulmonata: Charopidae). Memoirs of the Queensland Museum 34:11—20. SUSSMAN, R. W., G. M. GREEN & L. K. Sussman. 1994. Satellite imagery, human ecology, anthropology, and deforestation in Madagascar. Human Ecology 22:333-354. VAUGHT, K. C. 1989. A Classification of the Living Mollusca. American Malacologists Inc.: Melbourne, Florida. 189 pp. THE VELIGER © CMS, Inc., 2000 The Veliger 43(3):265—276 (July 3, 2000) Exploration of Morphospace Using Procrustes Analysis in Statoliths of Cuttlefish and Squid (Cephalopoda: Decabrachia)—Evolutionary Aspects of Form Disparity JEAN-LOUIS DOMMERGUES AnD PASCAL NEIGE Centre des Sciences de la Terre de 1’ Université de Bourgogne & UMR CNRS 5561, 6 Boulevard Gabriel, F-21000 Dijon, France AND SIGURD v. BOLETZKY Observatoire Océanologique de Banyuls, Laboratoire Arago, UMRCNRS 7628, F-66651 Banyuls-sur-Mer, France Abstract. This paper reports on a pilot study using, for the first time, a Procrustes type analysis of shape in exploring the morphospace of cephalopod statoliths. A total of 12 species of cuttlefish and squid (Decabrachia) from the Medi- terranean were analyzed, based on 18 homologous points (landmarks) chosen on the anterior statolith surface. For two species (one cuttlefish, one loliginid squid) size ranges were sufficiently large to reveal ontogenetic trends in statolith. Comparisons between species resulted in four well-defined sets of statolith morphology corresponding, respectively, to (1) sepiid cuttlefish, (2) Rossia (a large sepiolid squid), (3) myopsid squids, (4) oegopsid squids and small sepiolids. The morphological ‘‘dissociation’’ of large and small sepiolids suggests a relation between statolith size and shape ‘distinctness,’ and draws attention to the possibly paedomorphic shapes at the lower end of the size scale. INTRODUCTION Our study reconsiders the conclusion of Clarke & Mad- dock (1988b) as to phylogenetic information provided by Statoliths are calcareous particles that are attached to an statolith morphology, in (1) exploring morphospace pat- epithelial receptor complex inside the paired statocysts of terns in statoliths of several decabrachian groups, consid- cephalopods. They grow by periodic addition of aragonite ering variability at different levels from intraspecific to layers crystallized from the statocyst fluid. They are part intergroup (family) level, (2) assessing biological form of the macula/statolith system for the detection of gravity disparity between statoliths in terms of morphological and other linear acceleration. This gravity receptor system distances that are testable against the phylogenetic trees functions along with a complementary angular accelera- derived from molecular methods, and (3) identifying like- tion receptor system, which occupies the greater part of ly homoplasies by comparing traditional taxonomy, ex- the statocyst (Budelmann et al., 1997). The statoliths have ploitation of molecular patterns and statolith morphology. been studied intensively since the beginning of the 1960’s This approach should allow us to consider two evolution- (e.g., Young, 1960; Clarke & Maul, 1962; Lombarte et ary aspects complementary to one another, namely adap- al., 1997). In recent years, much attention has been given tive significance versus genetic fixation of statolith mor- to the growth layers observed in squid statoliths as po- phology. tential age markers (Jereb et al., 1991; Arkhipkin & Bi- Form disparity needs to be described in morphological zikov, 1997; Bizikov & Arkhipkin, 1997). terms applied to the disparity definition proposed by Raff Clarke & Maddock (1988a, b) came to the conclusion (1996), according to which disparity is the measure of that the shape of the statoliths depends little on their func- how fundamentally different organisms are. Methods of tion and heavily on the phylogeny. In contrast, the overall geometric morphometry (Procrustes analysis) have structure of the statocysts apparently reflects rather proved interesting on theoretical grounds (Bookstein, strongly the respective life style and locomotor activity 1991) and efficient in their applications to various zoo- of a species (Young, 1988, 1989). Thus the question re- logical groups (e.g., Tabachnik & Bookstein, 1990; Laur- mains open whether statolith form, even if it appears to in et al., 1994; Neige & Dommergues, 1995; David & depend little on the function of the statolith (Clarke & Laurin, 1996; Neige & Boletzky, 1997). These methods Maddock, 1988b), might nevertheless reflect some phys- are based on the utilization of anatomically conspicuous ical constraints related to the movement of the endolymph points (landmarks sensu Sneath, 1967; Bookstein et al., inside the statocyst. 1985; Bookstein, 1991). A given set of landmarks serves Page 266 The Veliger, Vol. 43, No. 3 Table 1 Traditional classification of the studied taxa (after Mangold & Portmann, 1989), and number of individuals (n) used for analysis. Order Suborder Family Subfamily n SEPIOIDEA Sepiidae Sepia officinalis Linnaeus, 1758 11 Sepia elegans d’Orbigny, 1835 11 Sepia orbignyana Férussac, 1826 11 Sepiolidae Rossiinae Rossia macrosoma (Delle Chiaje, 1829) 03 Sepiolinae Sepietta neglecta Naef, 1916 OS Sepietta oweniana (Pfeffer, 1908) 07 Sepiola sp. Ol TEUTHOIDEA Myopsida Loliginidae Loligo vulgaris Lamarck, 1798 32 Alloteuthis media (Linnaeus, 1758) 31 Oegopsida Enoploteuthidae Abralia veranyi (Riippell, 1844) Ol Ommastrephidae Ilicinae Illex coindetii (Vérany, 1837) 09 Todaropsis eblanae (Ball, 1844) 02 as a morphological descriptor. Comparison of the relative positions of these landmarks warrants localization and quantification of morphological differences between on- togenetic stages, individuals, or taxa. Such a representa- tion is called a morphospace (Neige et al., 1997); here it is given in the form of a phenogram for cephalopod stato- liths. MATERIALS AnD METHODS Our analysis covers nine genera: four Sepioidea, five Teu- thoidea (Table 1). This sample represents a wide range of decabrachian diversity and covers the greater part of taxa studied recently with molecular methods (Bonnaud et al., 1996, 1997; Boucher-Rodoni & Bonnaud, 1996). All the specimens were caught in the area of Banyuls-sur-Mer (western Mediterranean). Landmarks Although the statoliths of decabrachian cephalopods show a wide variety of forms, their structure is sufficient- ly constant to permit recognition of homologies. In our study, homologies are derived from a subdivision of the whole statolith in four basic compartments according to the terminology of Clarke (1978): attachment area or wing, ventral rostrum, lateral dome, and dorsal dome (Figure 1A). Nevertheless our approach of morphology using landmarks is different from the one chosen by Clarke & Maddock (1988b) who took linear measure- ments for subsequent processing by multivariate analysis. In this study, only the anterior side of the statolith is analyzed. Apart from the fact that the attachment area, which is one of the most important structural elements, is entirely exposed only on the anterior side, this side has the advantage of showing a clear partitioning, which al- lows one to recognize a large number of homologous points (Figure 1B). Since the position of maximum sur- face curvature in the lateral dome (Point 17) is used, our descriptor also provides some information on the relief of the anterior face. Figure 1C gives the localization and assessment of the 18 landmarks as adopted in this study, taking into account the terminology of Bookstein (1991). Data Acquisition and Morphometric Treatment Homologous points, localized on camera lucida draw- ings of statoliths, were seized using a digitizer (3SPACE® DIGITIZER). The adjustments permitting form comparisons were achieved with the LSTRA algo- rithm of the Procrustes program (David & Laurin, 1992). The phenetic trees derived from a distance matrix, cal- culated with the Procrustes program (see Appendix 1, 2, 3), were obtained using the Fitch algorithm of the PHY- LIP program (Felsenstein, 1990). The first step of the analysis was to compare all the individuals in pairs (Pairwise analysis of Procrustes soft- ware) within each species. In some cases, this step could allow one to surmise an ontogenetic trend in statolith morphology. To test the veracity of this observation, a phenetic tree is established that summarizes all the pair- wise comparisons. If such a trend is confirmed, the sam- ple is represented for the following interspecific analysis by two categories: one for small specimens, the other for large ones. To compare species, an average individual is calculated (Consensus Analysis option of Procrustes software) from all individuals included in the species, except in ontoge- netically marked species where two average individuals are calculated: one for small specimens, the other for large ones. RESULTS Variation within Species For Loligo vulgaris, if the largest individual is com- pared with the smallest, the adult statolith appears rela- J.-L. Dommergues et al., 2000 Page 267 A Lateral dome Dorsal dome Ventral rostrum onan oe hm C Number Type Description ' WwW W RR i i i ONNDNBPWNrK TU WHANANHA HWY NNNYMWNWNYWNWNNNYK VY NY LY Attachment area or Wing : Wing fissure : Wing shelf : Wing ventral indentation : Main spur : Wing dorsal indentation Medialmost point of wing spur Medialmost point of wing fissure Dorsalmost point of wing shelf Ventralmost point of wing shelf Medial contact of ventral rostrum and wing Lateralmost point of wing ventral indentation Medialmost point of main spur Lateralmost point of wing dorsal indentation Intersection of a parallel to the axis (5-10) and rostrum medial edge Ventralmost point of ventral rostrum Intersection of a parallel to the axis (10-12) and ventral rostrum Lateral contact of lateral dome and ventral rostrum Intersection of a parallel to the axis (12-14) and lateral dome Lateralmost point of lateral dome Intersection of a parallel to the axis (14-18) and lateral dome Medialmost point of lateral dome Maximum swelling of dorsal dome Dorsalmost point of lateral dome Figure 1 A. Statolith morphology. B. Localization of homologous points used. C. Definitions and respective types (sensu Bookstein, 1991) of the 18 homologous points used. tively wider (Figure 2). The tree constructed from the distance matrix (Appendix 1) is arranged from the small- est to the largest individuals (Figure 3). Thus the feeble morphological modification during growth is indeed a di- rected one. The same obtains in Sepia officinalis. In our sample, there is progressive shortening of the rostrum and widening of the attachment area (Appendix 2, Figure 4). Thus for the interspecific analysis Loligo vulgaris and Sepia officinalis are represented by two categories each: SMALL (smaller than 120 mm ML in L. vulgaris, smaller than 130 mm ML in S. officinalis) versus LARGE. The statoliths of other species analyzed under identical conditions (Sepia elegans, S. orbignyana, Alloteuthis me- dia) do not reveal growth-related morphological modifi- cations. Rossia macrosoma, Sepietta neglecta, S. oweni- ana, Sepiola sp., Abralia veranyi, Todaropsis eblanae, Illex coindetii are not represented by sufficiently different ranges of sizes to allow an ontogenetic analysis. Never- The Veliger, Vol. 43, No. 3 14 / /17 Bf 2f 16 # a7] 3 13. 64 74 Original species 12" 5 x Target species an <9 10 Loligo vulgaris —> Loligo vulgaris (ML = 42 mm) (ML = 235 mm) Target species Original species Figure 2 An example of comparison using LSTRA (comparison between two individuals of Loligo vulgaris of different sizes). A. Vector field resulting from Procrustes adjustment. B. Presentation used in this analysis. theless, for //lex coindetii, recent data from a very wide range of sizes reveal an ontogenetic change of statolith shape (Gonzalez & Guerra, 1997). Our present data cor- respond to stage 2 described by these authors. Even when sexes were distinguishable in sufficiently large numbers of individuals (Sepia elegans, S. orbigny- ana, Illex coindetti), no sexual dimorphism was detect- C) 235 Size classes (mm) C) 198 @[42 Sepia officinalis (ML = 48 mm) (ML = 191 mm) Page 269 Size classes (mm) @ [48 < ML < 83] © [84A? of 3.73 between LARGE Loligo vulgaris and Illex coindetii: Figure 5D, Appendix 3). In compar- ison with L. vulgaris, I. coindetii has statoliths in which the rostrum is markedly reduced, the lateral dome trun- cated laterally, but well developed dorsally and ventrally, and which has a longer attachment area. The three species of Sepia Linnaeus, 1758, have very similar statoliths (max. A? = 1.53, Appendix 3), always with a lateral dome forming a very distinct subspherical structure (which sets them apart from all the other taxa studied), a long and broad rostrum showing a truncated or broadly rounded end, and a large, massive attachment area (Figure 5E). The Sepiolinae form another group with high morphological coherence, although interspecies dis- tances are larger (max. LA? = 2.49, Appendix 3). Their statoliths are characterized by the lack of a ventral ros- trum (which implies superposition of points 5, 9, 10, 11; Figure 5F), a strongly (especially dorsally) reduced at- tachment area, and by a well-developed dorsal dome, which is only poorly demarcated from the lateral dome, however. The overall outline is slender in the statoliths of the Sepiolinae. The analyses reveal particularly marked differences be- tween Sepiidae and Sepiolinae (max. 2A? = 5.05, Ap- pendix 3). An intermediary position between these two sets is held by the sepiolid Rossia macrosoma (Figure 5G, H), which appears closer to the sepiids (max. YA? = 3.02, Appendix 3) than to the Sepiolinae (max. LA? = 4.43, Appendix 3). Apart from the lack of a (typically sepiid) subspherical lateral dome, the main difference between Rossia macrosoma and sepiid statoliths is the smaller ros- trum with a more rounded end and a wider lateral dome. Moreover, the pairwise analyses performed between Sepia elegans and Rossia macrosoma, on the one hand (see Figure 5G), and between Loligo vulgaris and Rossia macrosoma, on the other (Figure 5]), highlight the inter- mediary position of R. macrosoma between the Sepiidae and myopsid Teuthoidea. In contrast, the Sepiolinae ap- pear closer to the oegopsid Teuthoidea, as shown by the comparison between Illex coindetii and Sepietta oweni- ana (Figure 5J). DISCUSSION The phenetic tree derived from the distance matrix (Ap- pendix 3) gives a quantitative representation of relations Page 270 Loligo vulgaris (Small) —> Alloteuthis media AD = yELS Loligo vulgaris (Large) —> Illex coindetii SY A2 = 3.73 Rossia macrosoma Illex coindetii —> J Sepietta oweniana The Veliger, Vol. 43, No. 3 Loligo vulgaris (Large) —> B Alloteuthis media DYA2 = 1258 Sepia elegans —> FE Sepia officinalis (Large) DAZ = 1836 Rossia macrosoma —> H Sepietta oweniana AZ 3:81) Illex coindetti —> Todaropsis eblanae YA2 = 1.74 Sepietta oweniana —> F Sepietta neglecta YA2 = 1.31 Loligo vulgaris (Large) —> | Rossia macrosoma J.-L. Dommergues et al., 2000 Page 271 between statolith morphologies in the species studied (Figure 6). It reflects many of the taxonomic divisions traditionally accepted (Voss, 1977; Mangold & Portmann, 1989; Sweeney et al., 1992). For the Teuthoidea, the phenetic tree topology obtained with our method suggests a clear separation of Myopsida (with loliginid genera Loligo Lamarck, 1798, and Allo- teuthis, Wiilker, 1920) and Oegopsida (represented by two families, the Enoploteuthidae with Abralia Gray, 1849, and the Ommastrephidae with J//ex Steenstrup, 1880, and Todaropsis Girard, 1890). In contrast, the Se- pioidea do not cluster. Sepia and Rossia macrosoma, on the one hand, differ from the Sepiolinae (Sepiola Leach, 1817, Sepietta Naef, 1912), on the other. Thus the stato- liths of the Sepiolidae fall under two distinct morpholo- gies, which correspond to the Rossiinae and Sepiolinae, respectively. Clarke & Maddock (1988b:182) concluded that stato- liths largely confirm a broad pattern as expected from general systematic studies. However, for the Sepiolidae, they note that Heteroteuthis and Rossia are closer to the Sepiidae than is Sepiola. The morphological similarity between the statoliths of Rossia macrosoma and Sepia could be related to similar- ities in both animal and statolith sizes between Sepia and Rossia macrosoma, but the adult size of Heteroteuthis Gray, 1849, is closer to that of Sepiola or Sepietta. This has to be kept in mind in attempts to interpret the dis- parity between the statolith morphologies of Rossiinae and Sepiolinae in relation to functional constraints. Like- wise, the morphological similarity between statoliths of Sepiolinae and of oegopsid squids could be related to similarities in statolith size, but it cannot be related to the adult sizes of the animals, which are very different. Thus the question arises whether this similarity of statolith morphology might reflect a close phylogenetic relation- ship. This has to be discussed in relation to other pub- lished data, and especially with a close look at Idiosepius Steenstrup, 1881. Various data now call for a redefinition (or abandon- ment) of the higher taxa Sepioidea and Teuthoidea. With- in the Teuthoidea, the suggested transfer of Chtenopteryx Appelléf, 1890, from the Oegopsida to the Myopsida (to join the loliginds, as suggested by Young, 1991, and Brierley et al., 1996), is a minor change, but it may fore- shadow greater rearrangements. The removal from the Sepioidea, either of the Sepiolidae together with the Se- piadariidae and the Idiosepiidae (Fioroni, 1981), or of the Sepiolidae with the Idiosepiidae only (Clarke, 1988), has already launched a reassessment of sepiolid relationships (Boletzky, 1995). The resulting systematic changes within the former Sepioidea, or within the Decabrachia as a whole, are now highlighted by the possible transfer of Idiosepius to the Oegopsida based on nucleotide and ami- no acid sequences processed with the Neighbour Joining method, Bonnaud et al. (1997). Provided that this new position of /diosepius is not due to an artefact of data processing (especially relating to use of a distance method), the similarity of the statoliths of Idiosepius (Jackson, 1988) with those of Sepiolinae suddenly appears in new light. Indeed these statoliths are similar not only to those of Sepiolinae, but also to those of ommastrephid and enoploteuthid squids. Of course, this observation should not be taken to mean that the Sepiolinae or the whole family Sepiolidae would now have to follow in the wake of /diosepius in view of the other similarities (cf. Fioroni, 1981; Clarke, 1988). In contrast to the new position of /diosepius suggested by the above-mentioned molecular study, phylogenetic relationships of the Sepiolidae change depending on which subfamilies are included in the analysis: when rep- resented by Sepietta and Heteroteuthis the Sepiolidae cluster with the Sepiidae (Bonnaud et al., 1997), when represented by Rossia Owen, 1834, and Sepiola, they cluster with a group containing ommastrephids along with loliginids and sepiids, the latter two forming terminal sis- ter groups (Boucher-Rodoni & Bonnaud, 1996). An interesting parallel appears in the distribution of morphological characters in decabrachian spermatozoa. Healy (1989, 1990) observed that the respective forms in the cuttlefish Sepia, in the loliginids Loligo and Alloteu- this, and in the sepiolids Rossia and Sepietta are similar for the differentiation of a mitochondrial spur at the fla- gellar basis. In contrast, the acrosome comes in two forms, one (rounded) showing greater similarity between sepiid cuttlefish and loliginid squids, the other (elongate) showing greater similarity between different sepiolids (in- cluding Heteroteuthis). However, the spermatozoa of Het- eroteuthis are different from those of Rossia and Sepietta; they have a periflagellar mitochondrial sleeve instead of a mitochondrial spur. Surprisingly, this mitochondrial sleeve is very similar to what exists in Spirula Lamarck, 1799, whereas the (elongate) acrosomes of spirulid sper- matozoa are in their turn similar to those of the sepiolid spermatozoa in general. But other morphological and mo- lecular data do not suggest Spirula to be the closest rel- ative of sepiolids. In both instances, statolith morphology and spermato- zoan morphology, one has to cope with a mosaic of fea- tures. This situation necessitates a careful reassessment of Figure 5 Some examples of pairwise comparisons selected to illustrate the morphological differences within studied Deca- brachia. The Veliger, Vol. 43, No. 3 ® Sepia wd * a Sepia officinalis (Smal Sepia officinalis (Large) poe ate ti Sepia orbignyana Rossia macrosoma rN w Sepietta oweniana Sepioidea Sepiidae @ Sepiolidae (Rossiinae) «> ‘oli i, (Sepiolinae) — Teuthoidea (Myopsida) Loliginidae es (Oegopsida) Enoploteuthidae ia Ommastrephidae (Illicinae) WA Semi} — epiola sp. Gy Figure 6 Phenetic tree constructed from the quantification of differences and similarities in the statoliths of 12 analyzed species. All statoliths are drawn to the same scale. Note the overall size decrease from the top to the bottom of the figure. J.-L. Dommergues et al., 2000 Page 273 each of the characters considered. As to statolith mor- phology, the subglobular shape of the lateral wing in Se- pia then appears as a truly distinctive feature (a likely synapomorphy) that is much more significant as a phy- logenetic signature than the overall structure of the stato- lith. For the rest, it is the Jack of such distinctive features that leads to an ostensible mixture of systematic groups, similarity of form being inversely related to statolith size (Figure 6). More data from the subfamily Rossiinae, es- pecially from the small juvenile statoliths, are needed to see whether the statoliths of the Sepiolinae represent pae- domorphic forms derived from an ancestral situation clos- er to that of Rossia, or whether the latter results from hypermorphosis sensu Gould (1977). Complementary sets of ontogenetic data from the Heteroteuthinae might also be instructive. Of course, statoliths have to be viewed also on the background of statocyst morphology and related func- tional constraints. Describing the different statocysts in the three subfamilies of Sepiolidae, Young (1989:224) noted that the statocysts are basically all alike in being the shortest among all cephalopods relative to volume. Only in Rossia, some emphasis on the horizontal channel occurs (supposedly related to turning in the yawing plane). Discriminant analysis separates these statocysts from those of Sepia and places them close to those of the non-buoyant teuthoids. Perhaps this description of the statocysts provides some insight into the peculiar form of statoliths in Rossia; the well-developed lateral wings might reflect the emphasis on the horizontal channel and its supposed role in monitoring turning in the yawing plane. It also highlights similarities between sepiolids and non-buoyant teuthoids such as loliginids, ommastrephids, and enoploteuthids. 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The angular acceleration receptor system of diverse cephalopods. Philosophical Transactions of the Roy- al Society of London B 325:189—237. YOunNG, J. Z. 1991. Ctenopteryx the comb-fin squid is related to Loligo. Bulletin of Marine Science 49(1—2):148—161. Page 275 J.-L. Dommergues et al., 2000 eLl O¢l 6H CEL CEL C6 IL IGL SE? Oct brl BLL 80'l pel OL tL 6S I 08'c Ll’¢ Le? 9L’e 6e'L OL'L 00°¢ 99°L gQ'L OP'l €0'L Sel vet OL tL ev 9e"L ov kb 89° 00°¢ 8e'L 6c'l c0'?e 601 ge" OL pl t cel 8st 6v' kL Leb 6S'¢ 9b 90°¢ SLL 60'4 srt 860 6E'L 8S coe LO€ oe? coe Lye ZBL 60°C coe ele LO'?¢ pel Bel lo? QL'L v8'L LLL 260 6e'l 160 pot 69'L c0't LL’¢ 6L'e Sle plc 99°¢ cS €9'L Sel pl CASA LS'¢ c0'? 8r'C chem 6E'L QL’¢ Sel gc't bl t 90°¢ gle cL kt Aci LOL L6vL v8'b vel S6°¢ L €60 L LOL L 901 L Gob L 9E'L “L GGL | cok L 6c1 L 8b L L pel col LEL cGe ve? cel Spl 80¢ O0S¢ 96L E91 Lok TSTMNONONDONST COARTOHHOOH Sek ZL 96°0 Zeb byl LZ b Sl’¢ 6e'l Lvl Lyk 6S" oe? eo bk 9S°¢ pS'b 9e"L 90°L 6e'L OL? Ost pl 9S bk eo bk Sok Leb ov th LLL Bob ce? 60° elt LO0'¢ Aca Lvl 96° Sek OL? ZEL 99'L Zo‘ LLL 6S'L 6b verb Lvl br S6'L 9 6e°¢ eS Oot SEL lol cot OLE Eo? c0'L SSL 9p Os'L Loe 6v'L ov QL'L 9v't vel SLL Sle Sek pl t 680 Ze'l cL ht col Od"L 6b 8k°¢ vel Sv ev'k LoL 60°¢ Zyl Oe L Sel Ort Sle BLL LLe 9e"L ZL L Go"L cel 69'L OV’ 6EL SO'L OarannNnrroa SCOR CORCON CONN CO ILO Te rererrerrere ort Net srt ce Svl ev'l Oe'L Bk L 89'L 9e'L 9e'b Ov't ov vel OL gL lo? SO0°¢ 8c 1 LOL DEL LO'L 92 L cok ost cel c0'e 82h ZL 9r'L 88L Ze'l on kL OZ't 6EL Sc'k 89'1 66°14 eek Sv on ASE RG N oO Ba ed ed ao ed Oat cdi ool TOHOMHYtOT OyrOdDonrys On A ® ro werToMoaoruot SH SS SHS SU SSE rrrrrrrese Nr SB Ge —-reN so BLL LoL 660 co | S60 Oe L 601 9b Od'L Obl eel 8b 6e'L 9S"L Gel Lob vel pol LLé Gel evil ect 89°L eS'¢ 6r'L vse 89°L Ly 6LL c0'L vet bel tok 9 evil 68°0 Ov Oe 6b L cé kL 9p Oc'l Ono tToDoornr AN HDHAATH SS OLL# SOL# pOL# LOL#680# 880# L80# BZO# LZO# COOH EPO chO# LHOH LEOH OCO# 6cOH LcOH SZOH bcOH EcO# CcO#H |L2O# 6LOH SLOH ZLO# LLO# OLO# 600# SOO# cOO# LOO# ‘(SI9QUINU UOTJII[[OI :#) S1MD3]NA OS1]OT JO s[enplAIpUl ZTE IOJ (-VX) SOdMeU DOULISIG [ xtpuoddy LLL# OLL# SOL# VOL# LOL# 680# 880# Z80# 8Z0# LLO# cQ0# evo# cv0# LvO# LeO# OeO# 6c0# Lc0# ScO# vcO# €c0# ccO# LcO# 6L0# SLO# ZLO# LLO# OLO# 600# SOO# cOO# Page 276 The Veliger, Vol. 43, No. 3 Appendix 2 Distance matrices (2A?) for 11 individuals of Sepia offi- cinalis (#: collection numbers). #044 #046 #047 #048 #050 #112 #113 #114 #116 #117 #046 1.55 #047 1.36 1.20 #048 1.76 1.46 1.18 #050 1.90 1.92 1.23 1.20 #112 1.54 1.19 1.10 0.92 1.45 #113 1.28 1.30 1.20 1.43 1.82 1.07 #114 1.20 1.40 1.26 1.84 1.99 153 1.23 #116 156 0.89 1.09 1.38 1.68 1.15 1.10 1.46 #117 1.57 1.43 1.30 1.50 1.55 1.24 1.25 1.29 1.42 #134 1.28 1.46 0.95 1.37 1.39 1.31 1.37 1.16 1.40 1.05 Appendix 3 Distance matrices ({A*) for 12 decabrachian species studied. Amedi Icoin Seleg Sorby Sepsp LvulS Lvull SoffS SofflL Rmacr Snegl Sowen Todsp Icoin 3.43 Seleg 3.06 5.18 Sorby 3.42 5.25 1.27 Sepsp 4.43 4.16 4.72 5.05 LvulS 1.15 3.31 2.85 3.10 4.27 LvulL 1.58 3.73 2.11 2.44 4.31 1.01 SoffS 3.41 5.29 1.53 1.34 5.11 3.11 2.49 SoffL 3.63 5.49 1.36 0.98 4.82 3.33 2.68 1.11 Rmacr 3.01 4.43 2.28 2.74 4.43 2.56 2.39 3.02 2.9 Snegl 3.49 2.87 4.44 4.68 2.47 3.38 3.64 4.62 4.56 3.75 Sowen 3.11 2.52 4.55 4.78 2.49 3.08 3.41 4.67 4.67 3.87 1.31 Todsp 3.63 1.74 5.70 5.76 4.01 3.57 4.18 5.83 6.02 4.88 3.03 2.77 Avera 3.15 3.38 4.30 4.37 4.05 3.15 3.49 4.39 4.39 4.47 3.77 3.59 3.08 THE VELIGER © CMS, Inc., 2000 The Veliger 43(3):277—281 (July 3, 2000) Shell Size Variation and Aggregation Behavior of Littoraria flava (Gastropoda: Littorinidae) on a Southeastern Brazilian Shore PAULO R. S. MOUTINHO Instituto de Pesquisa Ambiental da Amazonia, IPAM, C.P. 6520, 66087-430 Belém, PA, Brazil AND CECILIA P. ALVES-COSTA Graduate Course in Ecology, Universidade Estadual de Campinas, C.P. 6109 13083-970 Campinas, SP, Brazil Abstract. Littoraria flava is a common snail on rocky shores and stems of mangrove trees along the southeastern coast of Brazil. However, studies on this species are absent. Our previous observation indicated a distance distribution pattern among shell size classes of this snail, depending on distance from the shore and occurrence in aggregation. We hypothesized that (1) snails with large shells would occur on tree stems far from the shore and (2) snails occurring in aggregation on rocky shores would be smaller than those occurring in isolation. We evaluated the effect of desiccation on the mortality of different size classes of snails experimentally submitted to a temperature of 40°C. We measured snail shells on 14 tree stems occurring in two mangrove strips (near and far from shore). Snails on tree stems of mangrove strips far from shore presented larger shells than snails near the shore. Snails in aggregations on rocky shores were smaller than isolated ones. The desiccation experiment showed that small snails (< 1 cm) lost water at a faster rate than larger ones (> 1.55 cm) and the mortality of small snails was above 70%. The distribution pattern of L. flava is probably related to differential shell size class tolerance to prolonged periods of emersion. Also, aggregation may be a behavioral mechanism to avoid water loss, as small snails are frequently found in aggregation and lose water at a faster rate. INTRODUCTION Shell size variation and aggregation behavior have been described for several marine gastropods (Feare, 1971; Butler, 1979; McQuaid, 1981; McCormack, 1982; Ver- meij, 1973) including several species of the family Lit- torinidae, which occur on rocky shores (Underwood, 1979; Chapman, 1994, 1995; Chapman & Underwood, 1996; Chapman, 1997). However, similar records for Neotropical littorinids are scarce (Britton, 1992; Maga- Ihaes, in press). In general, large shells may increase snails’ reserve of water and provide resistance to desiccation and high tem- peratures more than smaller shells. Thus large snails can inhabit higher zones of the rocky shores, reducing the risk of dislodgement by waves (Vermeij, 1973). Aggregations of snails may reduce the negative effects of desiccation and temperature (Garrity, 1984; Levings & Garrity, 1983), creating a moister microclimate. Aggre- gation may also allow snails to avoid displacement by waves (Boulding & Hay, 1993) or be the result of the presence of cavities or depressions, which are usually oc- cupied by snails (Chapman & Underwood, 1996). Cavi- ties and depression on rocky shores may protect the snails against physical displacement by waves, and retain mois- ture. In this work, we have described, for the first time for the Neotropics, the variation in the shell size and aggre- gation behavior of Littoraria flava King & Broderip, 1832, found in the zone among tides of rocky shores and on mangrove tree stems along the south coast of the State of Sao Paulo, southeast Brazil. We tested the following hypotheses: (1) snails with the largest shell lengths occur on the highest zone on the tree stems, whereas the smaller ones occur on the lowest zone, and (2) snails that occur on trees established in areas nearer to the tide line and at a lower elevation (i.e., those that are submerged first) are smaller then those occurring on distant trees at a higher elevation. Aggregations of L. flava are very common on shores along the southeast coast of SAo Paulo State. Our previ- ous observations suggested that the snails with smaller shells (< 1 cm in length) occupied the center of the ag- gregate, and those with very large shells (> 1.4 cm) re- mained isolated. We only observed aggregation during periods of low tide. We therefore tested the hypothesis that (3) snails occupying the center are, on average, smaller than those on the periphery, and that (4) the av- erage shell size of snails within the aggregates is smaller than the average shell size of isolated snails. Finally, we evaluated the effect of temperature on water loss and mortality of snails of different shell size classes in a con- trolled laboratory experiment. We expected that (5) small Page 278 snails, under high temperatures (> 40°C) would lose wa- ter at a faster rate than the larger snails, resulting in a higher mortality rate. MATERIALS anp METHODS Study Area This study was conducted at Araga Beach (23°45’S, 45°30'W), Sao Sebastiao, on the southern coast of the State of Sao Paulo, Brazil, during June 1997. The beach is sheltered from wave action, due to the presence of a small island known locally as Pernambuco Island. The distance between the island and the mainland is about 250 m with a topographical variation of 1.30 m. Contiguous to the island there is a small mangrove strip, formed mainly by the tree species Rhizophora mangle and La- guncularia racemosa. Another strip is situated near the continent, about 200 m from the first. Littoraria flava occurred on a rocky shore of the island and on the tree stems of the two mangrove strips. This study was carried out both on the island and along the mangrove strips, from 7—11 a.m. and with O—0.2 m of tide. Variation of Littoraria flava Shell Size on the Trees To evaluate the vertical variation in snail shell size on the mangrove trees (hypothesis 1), we measured shell length (spire length) and height of each snail on the tree stem. In addition, we compared the average shell length of the snails occupying tree stems between two mangrove strips (near and far from the tide line) (hypothesis 2). For each strip, we measured all the snails located on the stem of 14 trees and calculated a mean of the shell size for each tree between habitats. In order to test for a possible variation of shell shape (e.g., elongated or globular) as described for other species of littorinids (Chapman, 1995), we measured the difference in the relationship be- tween shell length and shell aperture for L. flava on trees and shore. Shell Size Variation in the Aggregates To test the hypothesis that the snails established in the center of aggregations are smaller than those occurring at the edges (hypothesis 3), we measured the shell length of snails in aggregations located on the rocky shore of Per- nambuco Island. We chose aggregates with circular form (12-20 snails/aggregates) that were established outside of rock cavities or depressions. We measured all of the cen- tral snails (those that were surrounded by other member of the aggregations on every side) and all of those at the edges (delimiting the aggregation). To evaluate whether the average size of the isolated snails was bigger than those in aggregations (hypothesis 4), we selected all isolated snails occurring in a radius of 50 cm around each aggregate. A snail was considered The Veliger, Vol. 43, No. 3 isolated if there was a minimum distance of 5 cm between it and any other snail. Statistical comparisons of all hypotheses were made with Student f-test. The assumption of normality was test- ed graphically, and homogeneity of the variances was checked with Bartlett’s test (Zar, 1984). Desiccation and Mortality of Different Size Class Snails under Higher Temperature To test the hypothesis that small snails (< 1 cm) suffer higher desiccation and mortality rates than the interme- diate (1.30—1.45 cm) and large sizes (> 1.55 cm), we subjected 10 snails from each size class to a temperature of 40°C inside a stove. The exposure period was 4 hours. In order to keep each snail isolated during the experi- ments, we enveloped each one with a pierced cotton tis- sue (tulle). We weighed all snails before the beginning of the experiment and after its completion. The difference between the weight of the two measurements allowed us to estimate the water loss suffered by the snails and ex- press it as a percentage. After the end of each experiment, we noted the number of dead snails in each size class to evaluate the relationship between the mortality and the shell size. The snails were put on the center of Petri dish- es wet with seawater, and we considered dead those that did not show any movement after a period of 12 hours. Voucher specimens were deposited in the Natural His- tory Museum of the Universidade Estadual de Campinas, Sao Paulo, Brazil. RESULTS Littoraria flava were abundant on both the mangrove trees and the rocky island shore. The aggregates were common on the rocky shore, but absent on the tree stems. Field observations suggest that the aggregation was max- imal during the low tide and especially when there was direct sun over the shore. The temperatures of the rock surface and of the air were around 24—26°C in the early morning (7—8 a.m.), but were higher than 30°C by mid- day. On rainy or cloudy days the aggregates on the rocky shore were smaller and less frequent. The relationship between the shell aperture and its length indicated that L. flava did not show variation in shape between snails occurring on the rocky shore and on tree stems of different mangrove strips (Figure 1). The average size of 778 snails sampled was 1.24 + 0.21 cm (mean + SD, range: 0.63—1.88 cm). The size of the snails’ shells did not vary as a function of their height on tree stems (hypothesis 1) (r = 0.11, p > 0.05, n = 397). On the other hand, the snails occurring on man- grove trees distant from the tide line and therefore pos- sibly subjected to a more extended emersion period (hy- pothesis 2), were bigger (1.30 + 0.09) compared to those on the trees nearer to the tide line (1.19 + 0.09, t = —3.02; n = 14; p < 0.01). P.R.S. Moutinho & C.P. Alves-Costa SHELL APERTURE (cm) 05 1.0 1.5 2.0 SHELL LENGTH (CM) Figure | Relationship between shell aperture and shell length in snails (n = 778), Littoraria flava, occurring on rocky shore (<) and on tree stems of two mangrove strips near (+) and far (©) from shore in Araga beach, Sao Sebastiao, Sao Paulo State, Brazil. Snails occupying the center of the aggregations were smaller (1.15 + 0.08 cm) than those at the edges (1.25 + 0.05 cm, n = 11, t = 3.4, p < 0.01) (hypothesis 3). Snails in aggregations were smaller (1.21 + 0.05 cm, n = 11) than isolated snails (1.30 + 0.05 cm, n = 6, t = —2.9, p < 0.01) (hypothesis 4). The experiment on the effect of temperature on the snails’ desiccation indicated an inverse relationship be- tween shell length and the percentage of water loss (Fig- ure 2). After 4 hours in the stove, small snails (<1 cm) lost 40% to 60% of their original water content (before desiccating), and had a mortality rate over 70%. The me- dium-sized snails (1.30—1.45 cm) lost 5% to 35% of wa- ter, and had a mortality rate of less than 10%. This also held true for the large snails (> 1.50 cm) that lost less than 10% of water (Figure 2). DISCUSSION A vertical gradient in the distribution of the sizes can often be detected in a shore, with larger snails occupying higher zones, and smaller ones the lower zones (Under- wood, 1979). Apparently, larger snails are better able to endure the stress resulting from prolonged periods of emersion (Vermeij, 1973; McMahon, 1990). On the other hand, variation in size can also be the result of differences in habitat conditions or the presence of cavities and other organisms. For instance, Chapman (1994) found that size of the shell in Littorina unifasciata is a function of its Page 279 50 40 ++ +4 5 ss * SMALL SNAIL w yy 30 o) i = 20 = < = 10 LARGE SNAIL 0) 0.5 1.0 1.5 2.0 SHELL LENGTH (CM) Figure 2 Relationship between water loss and shell length in snails Lit- toraria flava submitted to desiccation in a stove under 40°C, during 4 hours. occurence on the shore possibly determined by rock in- clination or the presence of cavities on its surface as well as the presence of barnacles. In this study, we did not verify the variation in the size of L. flava on the rocky shore. We judged this shore too narrow to produce a sharp gradient of shell size distribution as a function of the height of the shore. However, we could evaluate the ex- istence of this gradient by relating the size with the height of the snails on the mangrove trees and also by comparing the size of the snails between the two strips of mangroves. The absence of a relationship between shell size and the height of the occurrence of the snails on the stems, however, suggests that some conditions (moisture content along the stem, availability of food, and well-protected habitat) may serve to protect the snails from physical fac- tors. Desiccation, high temperature, and wind normally promote spatial segregation of snails by different shell sizes on the rocky shores (Vermeij, 1973). On the other hand, there was a sharp difference in the average size of the snails between the two mangrove strips, suggesting that there is a gradient in shell size related to wave ex- posure, as on the rocky shores (Chapman, 1994). The larger snails occupied the mangrove strip distant from the tide line where the potential desiccation is higher due to long periods of exposure. L. flava showed aggregation behavior only on the rocky shore, and not on tree stems. The aggregates can potentially confer advantages against predation and par- asitism, and help snails to avoid dislodgement by waves and reduce the stress provoked by desiccation and high The Veliger, Vol. 43, No. 3 temperatures (Chapman & Underwood, 1996 and cited references). Normally, littorinids aggregate during low tides (Chapman, 1997) and occur in crevices in the rocks (Chapman & Underwood, 1996). In this study, the same trend was observed for L. flava. Although not quantified, the aggregation pattern seemed less intense during rain events and overcast conditions, which indicates the influ- ence of temperature and desiccation as determinants for aggregation. Also, the effect of temperature and desic- cation can determine patterns of aggregation and the spa- tial distribution of the shell size in L. flava in the aggre- gates, with large ones occurring isolated, and smaller ones occupying the center of the aggregates. Although the physiological tolerance to desiccation and high tem- peratures can differ among snails on the same shore, these factors are apparently more harmful to smaller snails than to larger ones, simply as a consequence of the ratio be- tween area and volume of the shell, large for smaller snails. The experiments in a temperature-controlled stove confirm this hypothesis (4). The small snails were more susceptible to water loss and suffered higher mortality than the larger ones (Figure 2). Chapman & Underwood (1996) reported that L. unifasciata snails inside the ag- gregates held a greater quantity of water compared to isolated ones and that the aggregation was greater when the rocky substratum was dried. The size differences among the snails in the aggregates and the solitary ones found in this study suggest that the aggregation in L. fla- va, unlike that of L. unifasciata (Chapman & Underwood, 1996), is a behavioral response that reduces the stress induced by desiccation and temperature. Thus this behav- ior has a direct relationship with the size of the snails. This would explain, in part, the absence of aggregation in the snails that occur on tree stems, a substratum with supposedly higher humidity than the shore. Although the evidence suggests that temperature and desiccation can be primary factors in promoting aggre- gation, especially among the smaller snails, it is possible that the size segregation among the snails is the result of the higher capability of dislocation of larger snails. Garr- ity & Levings (1984) observed that the activity period of the mollusk Nerita scabricosta during low tide was in- versely related to the individual’s shell length. The same was found for Tegula funebralis (Marchetti & Geller, 1987). Therefore, small snails cover shorter distances than larger ones and return first to crevices to form ag- gregations. Furthermore, smaller snails can occupy small gaps in rock crevices and among the larger snails, facil- itating their occurrence in the aggregates. In any case, the aggregations may provide shelter to small snails, protect- ing those more vulnerable individuals from variations in physical conditions. Apart from the supposed advantages given by the formation of the aggregates, some field ob- servations suggested an extra advantage of aggregation to smaller snails. In the field and in the lab (we packed to- gether small and large snails in a glass pot), they were observed attached to the shells of larger snails and in continuous feeding activity. We sometimes found larger snails with the shells apparently “‘scraped,”’ and in some, the layer of microalgae that normally covers the large shells of L. flava totally removed by the smaller snails. Once each shell became completely ‘“‘clean,’’ smaller snails were not observed on them anymore. Therefore, it is possible that the aggregates also provide an extra food resource available on the shells of larger snails. Use of the shells of larger snails as feeding substratum by the smaller ones may explain in part the maintenance of ag- gregates in L. flava and possibly in other littorinid spe- cies. Acknowledgments. We are grateful to Dr. Luiz EK Lembo Duarte and to Claudia Alves de Magalhaes for their countless and valu- able suggestions during the course of this work and to Marcel Tanaka, Elizabeth Belk, Claudia Azevedo-Ramos, and Daniel Nepstad for revision of the manuscript. Support to P. Moutinho and C. P. Alves-Costa was provided by CNPq. This work was developed as part of the activities of the Marine Ecology Course (June 1997) of the Graduate Program in Ecology of the Univ- ersidade Estadual de Campinas. LITERATURE CITED BOULDING, E. G. & T. K. Hay. 1993. Quantitative genetics of shell form of an intertidal snail: constraints on short-term response to selection. Evolution 47:576—-592. Britton, J. C. 1992. Evaporative water loss, behaviour during emersion and upper thermal tolerance limits in seven species of eulittoral-fringe Littorinidae (Mollusca: Gastropoda) from Jamaica. J. Grahame, P. J. Mill & D. G. Reid (eds.), Pro- ceedings of the Third International Symposium on Littorinid Biology, 69-83. BUTLER, A. J. 1979. Relationships between height on the shore and size distributions of Thais spp. (Gastropoda: Muricidae). Journal of Experimental Marine Biology and Ecology 41: 163-194. CHAPMAN, M. G. 1994. Small-scale patterns of distributions and size-structure of the intertidal littorinid Littorina unifasciata (Gastropoda: Litorinidae) in New South Wales. Australian Journal of Marine and Freshwater Research 45:635—652. CHAPMAN, M. G. 1995. Spatial patterns of shell shape of three species of co-existing littorinid snails in New South Wales, Australia. Journal of Molluscan Studies 61:141—162. CHAPMAN, M. G. 1997. Relationships between shell shape, water reserves, survival and growth of highshore littorinids under experimental conditions in New South Wales, Australia. Journal of Molluscan Studies 63:511—529. CHAPMAN, M. G. & A. J. UNDERWOOD. 1996. Influences of tidal conditions, temperature and desiccation on patterns of ag- gregation of the high-shore periwinkle, Littorina unifasciata, in New South Wales, Australia. Journal of Experimental Ma- rine Biology and Ecology 196:213—237. FEARE, C. J. 1971. 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The effects of aggre- gation and microhabitat on desiccation and body tempera- ture of the black turban snail, Tegula, funebralis (A. Adams, 1855). The Veliger 30:127-133. McCormack, S M D. 1982. The maintenance of shore-level size gradients in an intertidal snail (Littorina sitkana). Oecologia. 54:177-183. Page 281 McMauon, R. F 1990. Thermal tolerance, evaporative water loss, air-water oxygen consumption and zonation of intertid- al prosobranchs: a new synthesis. Hydrobiologia 193:241— 260. McQualp, C. D. 1981. The establishment and maintenance of vertical size gradient in population of Littorina africana knysnaensis (Philippi) on an exposed rocky shore. Journal of Experimental Marine Biology and Ecology 54:77-90. UNDERWOOD, A. J. 1979. The ecology of intertidal gastropods. Advances in Marine Biology 16:111—210. VERMEU, G. J. 1973. Intraspecific shore-level size-gradient in in- tertidal molluscs. Ecology 53:693—700. ZAR, J. H. 1984. Biostatistical Analysis. Prentice Hall: Engle- wood Cliffs: New Jersey XIV + 718 pp. The Veliger 43(3):282-285 (July 3, 2000) THE VELIGER © CMS, Inc., 2000 NOTES, INFORMATION & NEWS New Information on a Poorly Known Late Paleocene Turrid Gastropod from Southern California and Vicinity Richard L. Squires Department of Geological Sciences, California State University, Northridge, California 91330-8266, USA Introduction Waring (1917), in his study of lower Tertiary marine mol- lusks from the Simi Hills and neighboring areas in Ven- tura County, southern California, described but did not illustrate the gastropod Bathytoma boundeyi Waring, 1917. Although this species is a turrid, it actually belongs in genus Parasyrinx Finlay, 1924. In addition, Waring’s species name is the senior synonym of another species name. The purposes of this report are to provide the first illustrations of the specimen Waring used to define his species and to provide a taxonomic update of this late Paleocene gastropod. Parasyrinx is interesting in that it is a predominantly Paleogene genus known only from the northeastern Pacific and New Zealand. As shown in this report, it appeared first in the northeastern Pacific but persisted longer in New Zea- land. The genus also has interesting paleotemperature and bathymetric distributions, first appearing in warm-water nearshore faunas, as shown in this report, but subsequently becoming typical of cool-water bathyal faunas. The following institutional acronyms are used: CAS, California Academy of Sciences, San Francisco; LAC- MIP, Natural History Museum of Los Angeles County, Invertebrate Paleontology Section, Los Angeles; LSJU, Leland Stanford Jr. University, Stanford (collections now housed at CAS); UCMP, University of California, Mu- seum of Paleontology, Berkeley; and UCR, University of California, Riverside. Systematic Paleontology Family TURRIDAE Swainson, 1840 Genus Parasyrinx s.1. Finlay, 1924 Types species: Pleurotoma alta Harris, 1897, by original designation; early Miocene, New Zealand. Parasyrinx boundeyi (Waring, 1917) (Figures 1—8) Bathytoma boundeyi Waring, 1917:81—82 (unfigured). Parasyrinx n. sp. Zinsmeister, 1983a: 70, pl. 4, fig. 27. Parasyrinx hickmani Zinsmeister, 1983b:1300, fig. 3 V, W; Zinsmeister & Paredes, 1988:13. Primary type material: CAS holotype 61901.01 [= LSJU holotype 193] of Bathytoma boundeyi. UCR holo- type 6668/36 and UCR paratype 4572/100 of Parasyrinx hickmani. Supplemental description: Shell small, up to 15.8 mm high (incomplete), pagodaform, five to six whorls, pleural angle about 50 degrees. Rounded angulation immediately posterior to suture; angulation with two moderately coarse spiral ribs and, on portion of abapertural side of body whorl of largest specimen, with several short and distinctly ophisthocline collabral nodes parallel to growth lines. Very fine spiral ribs on concave ramp of whorls; minutely beaded ribs near suture. Four very coarse spiral ribs just anterior to angulation on body whorl; ribs more closely spaced and finer anteriorly and slightly beaded on posterior part of neck. Growth lines on neck intersect very fine spiral ribs, forming a minutely cancellate pat- tern. Anal sinus U-shaped, in center of ramp. Inner lip smooth, with callus. Remarks: Waring (1917) reported a single specimen of Bathytoma boundeyi from his locality 4 [= CAS loc. 61901] in the upper Paleocene Santa Susana Formation on the south side of Simi Valley in the Simi Hills. This locality is in the lower part of the formation. The speci- men of B. boundeyi does not have the morphological fea- tures that are diagnostic of genus Bathytoma Harris & Burrows, 1891. These features are biconical shape, gem- mulose spiral ribs, an anal sinus whose apex is on a pe- ripheral carina, and a siphonal fasciole (Powell, 1966; Davies & Eames, 1971). Comparison of “‘Bathytoma’’ boundeyi with other fos- sil turrids revealed that Waring’s species is conspecific with Parasyrinx hickmani Zinsmeister, 1983b. Only two specimens of P. hickmani were reported by Zinsmeister (1983b). The holotype of P. hickmani is from UCR lo- cality 6668, and the paratype is from UCR locality 4572. Both are from the lower part of the Santa Susana For- mation in Meier Canyon where there are numerous float boulders from the almost completely covered (Squires, 1997) so-called Martinez marine member of Nelson (1925). Illustrations (Figures 3—8) of both of these spec- imens are provided here for comparison, and show, for the first time, the apertural view of each and the lateral views of the holotype. The holotype of “‘Bathytoma”’ boundeyi (Figures 1, 2) and the paratype of P. hickmani (Figures 3, 4) are both about 12 mm in height and are indistinguishable. The holotype of P. hickmani (Figures Notes, Information & News Page 283 Figures 1—8 Parasyrinx boundeyi (Waring, 1917). Figures 1, 2. Holotype CAS 61901.01, CAS loc. 61901, height 12.1 mm, x4.2. Figure 1. Apertural view. Figure 2. Abapertural view. Figures 3, 4. Paratype UCR 4572/100 of Parasyrinx hickmani, height 12.4 mm, <4. Figure 3. Apertural view. Figure 4. Abapertural view. Figures 5-8. Holotype UCR 6668/36 of Parasyrinx hickmani, height 15.8 mm, 3.2. Figure 5. Apertural view. Figure 6. Left-lateral view, using low-angle illumination. Figure 7. Abapertural view, using low-angle illumination. Figure 8. Right-lateral (outer lip) view. All specimens coated with ammonium chloride. 5-8) is slightly larger and differs only in the presence of four, or possibly five short, collabral nodes on the angu- lation on a portion of the abapertural side of the body whorl (Figures 6, 7). These nodes, which are not very obvious except under low-angle illumination, are not pre- sent near the outer lip and do not appear to be a constant morphologic feature. Based on the available evidence, ap- parently they are unique to the holotype of P. hickmani. By definition, the angulation should be smooth on this genus (Finlay, 1924; Powell, 1966; Zinsmeister, 1983b). A search of the LACMIP collection revealed additional specimens of Parasyrinx boundeyi at LACMIP localities 22307 (one specimen) and 22330 (11 specimens). They are from the same area where the type specimens of P. hickmani were collected. Most of these additional speci- mens are badly weathered. The largest ones, 14 mm high (slightly incomplete), show no nodes on the angulation. The shape of the shell, the shape and location of the anal sinus, and the subdued ornamentation argue for the assignment of Waring’s species to genus Parasyrinx. This genus ranges from late Paleocene to late early Miocene, with reported species known only from the southwestern coast of North America and from New Zealand (Zins- meister, 1983b; Beu & Maxwell, 1990). On the south- western coast of North America, Parasyrinx ranges from late Paleocene in southern California and northern Baja California to the latest Eocene and earliest Oligocene (Galvinian Stage) in northwestern Oregon and Washing- ton (Hickman, 1976). In New Zealand, Parasyrinx ranges from late Eocene to late early Miocene. The two nominal subgenera of Parasyrinx are Para- syrinx s.s. and Lirasyrinx Powell, 1942, and they differ primarily in the shape of the protoconch (Powell, 1942). The protoconch of Parasyrinx s.s. consists of two round- ed whorls, whereas that of Lirasyrinx consists of four whorls, with the first two smooth, and the remaining two spirally lirate (Powell, 1942; Beu & Maxwell, 1990). Par- asyrinx §.8. has been reported only from upper Oligocene Page 284 and lower Miocene rocks in New Zealand, and Lirasyrinx has been reported only from upper Eocene to upper Oli- gocene rocks in New Zealand (Beu & Maxwell, 1990; Maxwell, 1992). Hickman (1976), in her work on the Galvinian Stage occurrences of Parasyrinx in Oregon and Washington, did not use subgeneric subdivisions because she believed that the protoconch differences that distin- guish Parasyrinx s.s. from Lirasyrinx are of questionable value. Beu & Maxwell (1990), however, regarded these Galvinian Stage species as belonging to Parasyrinx s.1. These species are Parasyrinx kincaidi (Weaver, 1916), P. dickersoni (Weaver, 1916), and P. delicata Hickman, 1976. As there are no protoconch characters available from the specimens of P. boundeyi that would allow them to be assigned to subgenus based on existing criteria, these specimens can only be assigned to Parasyrinx s.1. When compared to the Pacific Northwest and New Zealand spe- cies of genus Parasyrinx, the specimens of Parasyrinx boundeyi are most like those of Parasyrinx kincaidi from Oregon and Washington, especially in terms of the round- ed angulation. Parasyrinx boundeyi differs from P. kin- caidi by having a much wider spire and coarser spiral ornamentation. At the localities where Parasyrinx boundeyi has been found, there are megafossil assemblages of similar taxo- nomic composition (Waring, 1917:71, 72; Zinsmeister, 1983a b). This entire megafauna, which is dominated by mollusks, is indicative of nearshore-marine conditions (Zinsmeister, 1983a, b). All these localities, furthermore, are in Parker’s (1983) ‘‘eastern facies’’ of the Santa Su- sana Formation. Using microfossil data, he determined that this facies was deposited in a deep-marine (middle bathyal) setting, and he reported that the nearshore-ma- rine megafauna must be displaced. Specimens of Parasyrinx boundeyi found in the Paleo- cene Sepultura Formation in northern Baja California are also associated with numerous other nearshore and warm- water (tropical to subtropical) gastropods and bivalves (Zinsmeister & Paredes, 1988). Specimens of the various species of Parasyrinx s.1. found in the Pacific Northwest Galvianian Stage are associated with outer neritic to bathyal, cool-water mollusks (Hickman, 1976, 1980). Late Eocene specimens of Parasyrinx (Lirasyrinx) from New Zealand are associated with warm-water (sub- tropical) mollusks, whereas early Miocene specimens of Parasyrinx 8.8. are associated with temperate-water mol- lusks (Beu & Maxwell, 1990). Bathymetric data for the New Zealand specimens, unfortunately, are poorly known. It is apparent, therefore, that members of genus Para- syrinx 8.1. were initially shallow-marine, warm-water taxa but later became restricted to cooler environs and, in the Pacific Northwest, also restricted to deeper environs. Acknowledgments. Lindsey T. Groves (LACMIP) provided ac- cess to the collections and procured casts of various fossil-turrid The Veliger, Vol. 43, No. 3 gastropods. Jean DeMouthe (CAS) and Marilyn Kooser (UCR) loaned specimens. LouElla R. Saul (LACMIP) and James H. Mc- Lean (Natural History Museum of Los Angeles County, Mala- cology Section) provided key observations. Literature Cited Beu, A. G. & P. A. MAXWELL. 1990. Cenozoic Mollusca of New Zealand. New Zealand Geological Survey Paleontological Bulletin 58:1—518, pls. 1—57. Davies, A. M. & FE E. EAMES. 1971. Tertiary Faunas. Volume 1. The Composition of Tertiary Faunas. Revised by FE E. Eames. George Allen & Unwin: London. 571 pp. FINLAY, H. J. 1924. The molluscan fauna of Target Gully. Part 1. Transactions of New Zealand Institute 55:495—516. Harris, G. FE 1897. Catalogue of Tertiary Mollusca in the De- partment of Geology, British Museum (Natural History). Part 1. The Australasian Tertiary Mollusca. British Museum (Natural History): London. 407 pp., pls. 1-8. Harris, G. EK & H. W. Burrows. 1891. Eocene and Oligocene Beds of the Paris Basin. Geologists’ Association for 1891. London. 129 pp. HICKMAN, C. S. 1976. Bathyal gastropods of the family Turridae in the early Oligocene Keasey Formation in Oregon, with a review of some deep-water genera in the Paleogene of the eastern Pacific. Bulletins of American Paleontology 70(292): 1-119, pls. 1-7. HICKMAN, C. S. 1980. Paleogene marine gastropods of the Keas- ey Formation in Oregon. Bulletins of American Paleontol- ogy 78(310):1—112, pls. 1-10. MAXWELL, P. A. 1992. Eocene Mollusca from the vicinity of McCulloch’s Bridge, Waiho River, South Canterbury, New Zealand: paleoecology and systematics. New Zealand Geo- logical Survey Paleontological Bulletin 65:1—280, pls. 1-30. NELSON, R. N. 1925. A contribution to the paleontology of the Martinez Eocene of California. University of California Pub- lications, Bulletin of the Department of Geological Sciences 15(11):397—466, pls. 49-61. PARKER, J. D. 1983. Lower Paleocene to lower Eocene, nonmar- ine to deep-marine strata of the Simi Hills, Ventura County, California. Pp. 3—22 in R. L. Squires & M. V. Filewicz (eds.), Cenozoic Geology of the Simi Valley Area, Southern California. Pacific Section, Society of Economic Paleontol- ogists and Mineralogists, Book 35: Los Angeles, California. PowELL, A. W. B. 1942. The New Zealand Recent and fossil Mollusca of the family Turridae. With general notes on turrid nomenclature and systematics. Bulletin of the Auckland In- stitute and Museum 2:1—183, pls. 1-14. PowELL, A. W. B. 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and Museum 5: 1-184, pls. 1-23. Squires, R. L. 1997. Taxonomy and distribution of the buccinid gastropod Brachysphingus from uppermost Cretaceous and lower Cenozoic marine strata of the Pacific slope of North America. Journal of Paleontology 71(5):847—861, figs. 1-5. WarInG, C. A. 1917. Stratigraphic and faunal relations of the Martinez to the Chico and Tejon of southern California. Pro- ceedings of the California Academy of Sciences, Fourth Se- ries, 7(4):41—124, pls. 7-16. WEAVER, C. E. 1916. Tertiary faunal horizons of western Wash- ington. University of Washington Publications in Geology 1(1):1-67, pls. 1-5. ZINSMEISTER, W. J. 1983a. Late Paleocene (“‘Martinez provincial Stage’’) molluscan fauna from the Simi Hills, Ventura Coun- ty, California. Pp. 61—70, pls. 1-4 in R. L. Squires & M. V. Notes, Information & News Page 285 Filewicz (eds.), Cenozoic Geology of the Simi Valley Area, Southern California. Pacific Section, Society of Economic Paleontologists and Mineralogists, Book 35: Los Angeles, California. ZINSMEISTER, W. J. 1983b. New late Paleocene molluscs from the Simi Hills, Ventura County, California. Journal of Paleon- tology 57(6):1282—1303, figs. 1—4. ZINSMEISTER, W. J. & L. M. PAREDES-MesiA. 1988. Paleocene biogeography of the west coast of North America: a look at the molluscan fauna from Sepultura Formation, Mesa San Carlos, Baja California Norte. Pp. 9—22, pls. 1-3, in M. V. Filewicz & R. L. Squires (eds.), Paleogene Stratig- raphy, West Coast of North America. Pacific Section, So- ciety of Economic Paleontologists and Mineralogists, West Coast Paleogene Symposium Vol. 58: Los Angeles, Cali- fornia. Localities Cited Topographic base map is the U. S. Geological Survey, 7.5-minute, Calabasas Quadrangle, California, 1952 (pho- torevised 1967). CAS 61901 [= LSJU 2695 = UCMP 3776]. Just east of the letter “‘e’’ in Runkle Canyon. LACMIP 22307. Fine-gained gray concretionary sand- stone outcropping on west side of Meier Canyon in vicinity of the word “‘Canyon”’ in Meier Canyon. LACMIP 22330. Beds cropping out on nose of spur on west side of Meier Canyon, approximately 193 m north of second ‘“‘n” in Meier Canyon. UCR 4572. Concretion from west side of Meier Canyon, 549 m N23°W of hill 1658 in NW corner of quadrangle. UCR 6668. Concretion from west side of Meier Canyon, 396 m due west of hill 1658 in NW corner of quadrangle. International Commission on Zoological Nomenclature The new and extensively revised 4th Edition of the In- ternational Code of Zoological Nomenclature has now been published and is in effect from 1 January 2000. The price is US $65 or £40, but discounts are offered to in- dividuals buying the Code for personal use or to institu- tions buying five or more copies. Full details of how to buy copies are given on the Commission’s website (www.iczn.org) or may be obtained by e-mailing “iczn@nhm.ac.uk’’. The following Opinions concerning mollusks were published on 30 September 1999 in Volume 56, Part 3 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Exec- utive Secretary, LC.Z.N., % The Natural History Muse- um, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). Opinion 1939. Osilinus Philippi, 1847 and Austrocochlea Fischer, 1885 (Mollusca, Gastropoda): conserved by the designation of Trochus turbinatus Born, 1778 as the type species of Osilinus. Opinion 1931. Campeloma Rafinesque, 1819 (Mollusca, Gastropoda): conserved. Opinion 1932. Holospira Martens, 1860 (Mollusca, Gas- tropoda): Cylindrella goldfussi Menke, 1847 designat- ed as the type species. The Veliger 43(3):286—288 (July 3, 2000) THE VELIGER © CMS, Inc., 2000 BOOKS, PERIODICALS & PAMPHLETS Describing Species Described Describing Species; Practical Taxonomic Procedure for Biologists. By JUDITH E. WINSTON. 1999. Columbia University Press, New York. ISBN 0-231-06824-7 (hard- bound), ISBN 0-231-06825-5 (paperback). Although it is widely acknowledged that describing the biodiversity of the world is an enormous and urgent task, practical advice on this aspect of systematic biology is often lacking from life sciences curricula. Many biolo- gists complete their degrees with a dim awareness that some authority (“‘they’’—meaning, perhaps, the Interna- tional Commission on Zoological Nomenclature and the International Commission on Zoological Nomenclature and the International Botanical Congress) exerts gover- nance over the process, but with little concept of how it works. In fact, taxonomy is one of the greatest democracies in science. Cooperation with the rules in zoology and botany is voluntary, although mediated by the judgment of rep- utable journal reviewers and editors. For better or for worse, a published species description needs no commit- tee’s imprimatur to enter into the crucible of systematics. Formal nomenclature, an important tool of taxonomy and the subject of the International Code of Zoological No- menclature, is algorithmic (Lindberg, 1999). Like algo- rithms in general, it is substrate-neutral (the power of the procedure is due to its logical structure, not the causal powers of the materials used in any given instance, nor the person performing the operation); it consists of simple component steps (perhaps even ‘“‘[s]Jimple enough for a dutiful idiot to perform’? [Dennett, 1995:51]); and results are guaranteed. The result may not be exactly what the investigator wishes (e.g., a favorite name may lose out because it is a junior synonym), but there will be a result. Describing Species is intended to clarify taxonomic procedure (“the practical process of identifying, recog- nizing, researching, or redescribing a taxon for scientific publication according to the current rules of biological nomenclature;”’ p. 9) for the lay biologist. It aims to help non-taxonomists who find new species in the course of their research—and cannot, for reasons well foreshad- owed in the introduction, find a specialist willing or able to provide a timely description—to perform the necessary background studies and write publishable descriptions of those species. Finally, it is also a handbook to help the professional taxonomist improve his or her practice. For this last function, it is a welcome replacement to old but outdated standbys (e.g., Mayr et al., 1953; Blackwelder, 1967). The book is organized into four parts: (1) an introduc- tion to the history of species descriptions and the naming of organisms (with advice not to skip this part, because understanding this history helps make sense of the pro- cedure as practiced today); (2) the background research necessary before a species can be described; (3) the writ- ing of species descriptions and their publication; and (4) more advanced topics such as key construction, redescrip- tion and revision, and description of taxa of other ranks than species. In the first part, the author, who is Director of Research at the Virginia Museum of Natural History, an experi- enced invertebrate taxonomist, and a science historian, traces briefly the growth of formal nomenclature out of folk taxonomy. She points out that the type concept— fundamental to both the current botanical and zoological codes of nomenclature—could not have come about until after Darwinism had become established and the essen- tialist concept of species rejected. She tabulates the dif- ferences between the botanical and zoological codes. She touches on possible future codes unifying nomenclature for plants and animals and acknowledges the potential for phylogenetic classification systems. Finally, she points out the present and potentially much greater future impact of high-speed data processing and global connections via the Internet. Part Two, “‘Recognizing Species,”’ briefly reviews ma- jor species concepts and speciation phenomena and hints at the process by which a taxonomist judges a character to have taxonomic significance. It stresses the importance of literature search, with cautionary examples of authors who rushed into print without making sure that what they were describing was both new and real. It sets forth the components of a taxonomic description and how to read them. The place of museums in the process—their unique value and also their limitations—is described with the perspective of a professional in the industry. Like other sections, this one has an extensive and useful bibliogra- phy, including Internet resources. With Part Three, ‘“‘Writing Species Descriptions,”’ we get down to the nuts and bolts: the types of papers that may carry taxonomic descriptions (with examples by ti- tle); the component parts of a descriptive paper; headings and synonymies; etymology of scientific names; type and voucher material; diagnosis and description (two concepts not, in this editor’s experience, always correctly differ- entiated in the minds of publishing systematists); taxo- nomic discussion; ecology and distribution; material ex- amined; and not least, publication itself. To taxonomists with long experience in publishing, this may sound too much like a basic cookbook to be of interest; but I would Books, Periodicals & Pamphlets Page 287 encourage reading it as a refresher course, anyway. To potential authors of systematic papers for The Veliger, I particularly commend the advice on illustrating taxonom- ic descriptions (pp. 231—238). The fourth part addresses more specialized, but not less important, aspects of taxonomic practice: how should the treatment of subspecies differ from that of species; the traditionally neglected concept of the genus; special rules (and problems arising from the absence of formal rules) in the case of higher categories. Understandably, because Describing Species is a handbook, not a treatise, some philosophical aspects of these topics are merely pointed out or referenced. In a few cases, however, the practical advice is limited by a canonical, not phylogenetic, un- derstanding of systematics. For instance, the statement ‘““Genera are defined not by one character, but by a group of carefully chosen characters” (p. 341) is a thinly veiled statement of the magnitude-of-difference criterion that is common in traditional taxonomic thinking—that there must be more differences for something to qualify at a particular rank level. (To quickly see the fallacy of this, ask yourself how many uniquely derived character-states diagnose the higher taxon Gastropoda.) In phylogenetic taxonomy, it is the deployment of a character-state among the array of taxa studied (i.e., is it an apomorphy of a clade?) that determines whether that state is diagnostic (not “‘definitive’’) of a taxon. The advice on monotypic genera is similarly mislead- ing: ““‘when most of the taxonomic characters found in a new species do not correspond well with those in other species of any known genus and cannot, by a reasonable amount of modification of the generic diagnosis, be made to fit into any described genus, then a new genus should be created” (p. 341). But the proposal of a monotypic genus adds no new information to that included in the description of its type species. Basically, all the author of a monotypic genus is saying is, “this species is different and the differences impress me enough that I wish to recognize them at the level of genus.’’ Because no ob- jective criteria exist as to how different a taxon must be to qualify as a genus, such a decision is entirely subjec- tive. Conway Morris (1998:183) called it “‘really an eva- sion [that] solves nothing, at least in the context of evo- lution.”” He was referring to proposals of new phyla, but the principle holds equally true at lower taxonomic levels. Chapter 19, on keys and key construction, is seminal. Like diagnoses, keys distill the essence of a taxonomist’s observations and test his or her ability to communicate those observations—that is, to make them useful to the world at large. In this context, I was pleased to see reference to the DELTA (Descriptive Language for Taxonomy) system (Dallwitz, 1980; available at www.biodiversity.uno.edu/del- ta/), which is useful not only for generating keys but also as a taxonomist’s self-test of his or her own conceptions. Like other democracies, taxonomy functions best when its citizens are responsible and well informed. Describing Species is an important contribution toward that goal. I hope and trust that in addition it fulfills its aim to be a positive solution to the crisis of describing imperiled bio- diversity; that it improves the visibility of descriptive tax- onomy in biological curricula; and that it helps make life easier for biological editors everywhere. B. Roth Literature Cited BLACKWELDER, R. I. 1967. Taxonomy: A Text and Reference Book. New York: Wiley. 698 pp. Conway Morris, S. 1998. The Crucible of Creation. Oxford University Press: Oxford. xxiii + 242 pp. Da.tLwitz, M. J. 1980. A general system for coding taxonomic descriptions. Taxon 29:41—46. DENNETT, D. C. 1995. Darwin’s Dangerous Idea. Touchstone: New York. 586 pp. LINDBERG, D. R. 1999. [Review of] Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. 2nd ed. The Veliger 42(2):194—199. Mayr, E., E. G. LINsLEy & R. L. USINGER. 1953. Methods and Principles of Systematic Biology. McGraw-Hill: New York. 328 pp. A Field Guide to Marine Molluscs of Galapagos by CLEVELAND P. HICKMAN, JR. & YVES FINET. 1999. Sug- ar Spring Press, 802 Sunset Drive, Lexington, Virginia, USA. 150 pp. US $18.95. The benthic marine fauna of the Galapagos has long been a topic of interest to zoologists and biogeographers. This field guide, according to its authors, is intended to bridge the gap between the molluscan literature and ex- isting field guides, by describing and illustrating about one-quarter of the species of benthic marine mollusks found in the Galapagos: It certainly supplements the mol- luscan literature well, just by providing good illustrations, which have been lacking in many guides to the local fau- na. The guide covers 56 bivalve species, nearly 200 gas- tropod species, including heterobranchs, and four chiton species. The shell of each species is photographed, and a number of live animal photos are included, in the shelled taxa as well as in the shell-less groups. For many species, photos of uncleaned and juvenile shells are included as well, which will be very helpful to amateurs and natu- ralists who are likely never to see an animal with a clean or pristine shell. The one drawback here is that the pallial lines and sinuses of the bivalves are not shown for most bivalve species. The descriptions include Latin and com- mon names, size, descriptive information, habitat, and geographic range. The introductory sections of the guide provide a gen- eral map of the islands, a discussion of the origins and Page 288 endemicity of the Galapagos fauna, an introduction to no- menclature, and the usual discussion of the terminology used to describe molluscan shells. The introduction to no- menclature is better than the usual because it includes a discussion about the Law of Priority, with which non- specialists may not be familiar. The classification used is based on Keen (1971), with the addition of a few changes at the family level. The higher classification, however, has unfortunately not been updated to reflect recent advances in molluscan system- atics. This is doubly unfortunate because valid higher tax- on names could have been used very easily in this par- ticular guide without causing undue confusion to users who are unfamiliar with the continuing saga of molluscan systematic research. Species in this book are organized in families within the three major groups. The only one of the three groups that is subdivided further is Gastropoda, which is divided into prosobranchs, opisthobranchs, and pulmonates, all of which are either invalid or of ques- tionable validity. This level of subdivision is not included The Veliger, Vol. 43, No. 3 in the Table of Contents, although it probably should have been. Thus more appropriate higher taxon names could have been used in this book with little change in the order of species, and no change to the Table of Contents or to the ease of finding a given species. The back sections of the book include a short glossary, list of references and, index. The list of references is quite well rounded, including some popular guides as well as taxonomic references and also some biological and en- vironmental references. To summarize, this guide is attractive, reasonably com- plete, and will undoubtedly be very useful to scientists and naturalists researching or observing Galapagos mol- lusks. Marta J. deMaintenon Literature Cited KEEN, A. M. 1971. Sea Shells of Tropical West American: Ma- rine Mollusks from Baja California to Peru. 2nd ed. Stanford University Press: Stanford, California. xiv + 1064 pp. Information for Contributors Manuscripts Manuscripts must be typed, one side only, on A4 or equivalent (e.g., 842” X 11”) white paper, and double-spaced throughout, including references, figure legends, footnotes, and tables. All margins should be at least 25 mm wide. Text should be ragged right (i-e., not full justified). 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Miller. 1973. The Mollusks of the Arid Southwest. University of Arizona Press: Tuc- son. xvi + 271 pp. c) Composite works: Feder, H. M. 1980. Asteroidea: the sea stars. Pp. 117-135 in R. H. Morris, D. P. Abbott & E. C. Haderlie (eds.), Intertidal Invertebrates of California. Stanford Univer- sity Press: Stanford, Calif. Tables Tables must be numbered and each typed on a separate sheet. Each table should be headed by a brief legend. Avoid vertical rules. Figures and plates Figures must be carefully prepared and submitted ready for publication. Each should have a short legend, listed on a sheet following the literature cited. Text figures should be in black ink and completely lettered. Keep in mind page format and column size when designing figures. Photo- graphs for halftone reproduction must be of good quality, trimmed squarely, grouped as appropriate, and mounted on suitably heavy board. 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CONTENTS — Continued NOTES, INFORMATION & NEWS New information on a poorly known late Paleocene turrid gastropod from southern California and vicinity RICHARD Ls SQUIRES ise veces tale de ei c5ine Silo tel URiel ae ge epee Meare elas nee HOLS Re BOOKS; PERIODICALS 8 PAMPBHIEET So iecic 50 -schauswennniers) lene eee eee mA A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler (1901-2000), Founding Editor Volume 43 EXMTFSONTA oct 162000 5) QE | ISSN 0042-3211 October 2, 2000 Number 4 CONTENTS DrakudoltStohlen @901=2000)iMemoriallilssue! So Wee ee nected dsl ods es sre wie oie i Dr. Rudolf Stohler: some personal remembrances SAIN SHS ERIS Cheer hei Paani paint criia ae MMs Leib Wwe aealters wal Mi NVI UMW CoN Re vatldlotgs chassis il Distribution, reproduction, and shell growth of limpets in Port Valdez, Alaska ARN YER LAN CHARD AND ba OWARD >My BEDER Y= KS q ou > n fo) Z oO A. Blanchard & H. M. Feder, 2000 Table 3 Spearman rank correlations between monthly average counts of limpets at each tidal height sampled. The crit- ical level for significance testing is the Bonferroni cor- rected a* = 0.0056. Tectura Tectura Lottia persona fenestrata species Tectura fenestrata = (ORIN: — a Lottia species ns OS9** _- Juveniles —i(0) kOe 0.59** O35 * 0.0001 = P < 0.0083, ** P < 0.0001. pling at each site. Water samples for suspended sediment and salinity were collected at low tide from the shoreline. Salinity measurements were determined using a portable American Optical compensated salinity refractometer. In the laboratory, the water samples were filtered on pre- weighed 0.45 mm Millipore filters. The filters were dried to a constant weight at 60°C and suspended sediments calculated (mg 1"'). Transects were established at each site and sampled at permanently marked points separated by a 40 cm vertical distance. Intervals along the transect were marked start- ing from the upper edge of the lichen (Verrucaria spe- cies) zone in the high intertidal down to the lowest tidal height available in May 1988. The tidal heights (m) of each sampling location were calculated from predicted tide heights (NOAA tide tables). Four to eight 0.04 m? quadrants were sampled at each marked interval by a ran- dom toss of a 20 X 20 cm sampling frame onto the sub- strate. The number of limpets of each species observed in the frame was recorded. Within Port Valdez, some of the dominant limpets are cryptic so sampling included searching underneath rocks and crevices observed within the sampling frame. For abundance estimates, densities of Lottia pelta and L. borealis were recorded as Lottia species due to difficulties in separating the two species in the field. Additionally, no attempt was made to identify juvenile limpets (< 5 mm) to species. Since the abun- dance data were too numerous to present, plots of limpet abundance were made for two selected tidal height rang- es—0.35 to — 0.15 m CD and 2.05 to 2.25 m CD, rep- resenting the abundance of limpets at a high and low tidal range. However, the full data set was utilized for other analyses. Stereological methods were applied to assess the re- productive cycle of T. persona. Up to 12 limpets were Page 293 Table 4 Spearman rank correlations between monthly average counts of limpets by species and measured environmental variables. The critical level for significance testing is the Bonferroni corrected a* = 0.0031. Tectura perso- Tectura Lottia Juven- na fenestrata species iles Air Temperature ns 0.50** ns ns Water Temperature ns 0.68** ns ns Salinity ns — 0.64** ns ns Suspended Sediment ns ns ns ns * 0.0001 = P < 0.0083, ** P < 0.0001. processed for each site from every sampling period al- though numbers varied due to sampling difficulties. Lim- pets were preserved in Baker’s Formal-Calcium in the field and stored in 70% ethyl alcohol in the laboratory. Histological preparation of reproductive tissues followed procedures outlined by Lowe et al. (1982). Up to 20 fol- licles from a limpet were classified by stereological ob- servation, and the reproductive stage of an individual was assigned as the category containing the greatest count of follicles. The tissues were placed within the following categories adapted from Orton et al. (1956): Neuter (no identifiable reproductive tissues); Developing (small de- veloping reproductive tissues in which developing or- ange-colored eggs or cream-colored sperm were observ- able); Ripe (the gonad is thick and swollen; orange eggs are compressed and polygonal in shape and sperm fills the male follicles); Spawning (eggs round due to release of some gametes and male follicles show space in the center where sperm has been released); and Spent (only residual eggs or sperm are present). Limpets were sampled for growth and age measure- ments at every tidal height at the study sites in March and June 1990 using the quadrant-sampling method de- scribed above. Total length of limpets was measured (+ 0.01 mm) using a video image analysis unit (consisting of a video camera attached to a dissecting scope and to a digitizer pad for measurements). Limpets were aged by counting the major growth disruption rings on shells. The age of a limpet was taken to be the sum of the observed growth rings. Since movement of a limpet to the next age class would occur with the onset of new growth, an age- O limpet (limpets settling in winter/early spring) would have a shell composed of up to one season’s worth of Figure 2 Plots of environmental variables from the four sites in Port Valdez, Alaska. The plot labeled a is air temperature (C°), b is surface-water temperature (C°), c is salinity (%c), and d is suspended sediment (mg L~'). Page 294 The Veliger, Vol. 43, No. 4 Tectura fenestrata 2.05 to 2.25 m CD @ a P S O° N D J F M AM J- J A SS OPN Died EF Me AGMe ae Awa 1988 1989 1990 BEG Tectura persona 2.05 to 2.25 m CD Ss O ND S FM AYM I J A SO N= Dd SIM AS Meiers Ss 1988 1989 1990 Tectura persona -0.35 to -0.15 m CD e eatra St SO ON, D J F OMA Md id (Ae S| 3O) NG DJ) FE Ml ACUMa Je Jee Aas 1988 1989 1990 SECS 9 — Esso —_e—_ Cas 7M AB B4 SB Figure 3 Abundance (ind. m*) of Lottia spp., juveniles, Tectura fenestrata, and T. persona at four sites from Port Valdez, Alaska. Error bars are + 2 standard errors. A. Blanchard & H. M. Feder, 2000 Page 295 Lottia spp. 2.05 to 2.25 m CD o,.8, © A e S 0 N D JFM a M J J mi “Ss 0 IN DY J MM a M J ai A S§ 1988 1989 1990 ; Lottia spp. -U.35 to -0.15 m CD 50 400 300 200 Ind. m? 100 7 4 S -@ IN 1D) ghd WE ANIME Bl ah A TSO IN 1D ai eK Mt JI Jd A S 1988 1989 1990 Juveniles 2.05 to 2.35 m CD ond K+ MO OCH CCE Oth Hk (I OCC Onan one S re) N D J F M ‘A M J) ORAS SHORING Dino i NIG SAG Vit Jai Jit PAGS 1988 1989 1990 Juveniles -0.35 to -0.15 m CD 1988 1989 1990 Pas eats FE ERS ah ae cain 25s 7M AB B4 SB Figure 3 Continued. Page 296 The Veliger, Vol. 43, No. 4 7 Mile Beach 100 Y © = Y 80 = : = |ZS s 60 76 |BSpent 2 = @ Spawning ¢ 40 B Ripe : = | & Developing ae UG Nentersaa 0 100 » s 80 = a a 60 = | @ Spent pe = & @ Spawning geese = = | @ Ripe : 20 = = | 8 Developing ; =| 72 GNeuter 0 = = 1988 1989 1990 100 — a == a. =a: a 60 == Spent | — — | fe Ss & @ Spawning S28 =e = | | @Ripe ° = & = ; 5 20 == = | & Developing | ‘ | ‘ONeuter 0 | J M 1988 1989 Shoup Bay 100 2 g 80 Y, Z D : Pe ; = = | @ Spent 5. 60 = = | te = = @Spawning | are = = ®@ Ripe | 8 20 = = © Developing ee = = O Neuter 0 = a M Figure 4 Bar charts of percent reproductive stage of Tectura persona by month for the four sites in Port Valdez, Alaska. A. Blanchard & H. M. Feder, 2000 Table 5 Summary of regression analysis of estimated age and growth of Tectura persona from sites in Port Valdez. n = 812. Coeffi- t- cient SE statistic P-value Intercept 7M, SB 2.49 0.207 12.07. <0.0001 Slope 7M, B4, SB 1.63 0.038 42.34 <0.0001 Intercept AB 1.71 0.410 4.17 <0.0001 Intercept B4 0.36 0.154 2.31 0.0211 Slope AB — 0.31 0.073 — 4.22 <0.0001 growth with no obvious growth ring. Movement to sub- sequent age classes occurs with the onset of new growth. Although preliminary marking studies (Feder et al., 1992) were not entirely conclusive in determining whether rings on limpet shells represented annual growth, these rings were the best available estimate of limpet age. Regression analysis includes regression of limpet length against tidal height and estimated age for each site. Data Analyses The analyses of the abundance and reproductive data are primarily descriptive. The cryptic behavior of some species made accurate abundance estimates impossible, and abundance data are presented as means and standard errors. Nonparametric correlation analyses were per- formed to assess associations between environmental var- iables and limpet abundance with data from all sites com- bined. The significance level of the correlation analyses were corrected for the number of comparisons by the Bonferroni correction of a* = a/n where n = # compar- isons made. Growth data were analyzed using linear regression pro- cedures. Preliminary analysis of the growth to age rela- tionship for T. persona indicated an extremely shallow concave growth curve, and nonlinear regression estima- tion procedures could not adequately fit the von Berta- lanffy model, a normal model for limpet shell growth (Branch, 1981), to the data. Initial linear regression anal- yses indicated only slight deviations from the linear mod- el at the extremes of the growth measurements. Thus, growth is modeled using linear regression (Neter et al., 1990) as an approximation to the very shallow, concave growth curve of limpets in Port Valdez. The potential for bias in the regression coefficients due to errors in the estimation of age of limpets was recognized. Neverthe- less, the regression analyses are presented to provide a descriptive framework useful for future investigations. Data files pertaining to this study may be accessed on the World Wide Web at www.veliger.org. Page 297 RESULTS Seasonal trends in environmental conditions exhibited marked extremes (Figure 2). Air temperatures measured at the time of sampling ranged from — 6.7°C to 18°C and surface-water temperatures from 1.5°C to 13.0°C. Salinity ranged from 0% to 31%c and turbidity from 1 mg L™! to 98 mg L~'. ANOVA comparisons of the environmental variables indicated no significant differences (p < 0.05) in mean values between sites over the course of the study. Correlation analysis of environmental variables indicated significant positive correlations (p < 0.0083) between air and water temperatures, negative correlations for both air and water temperatures to salinity, a low positive corre- lation of suspended sediment with salinity, and a negative correlation of suspended sediment with air temperature (Table 1). Field observations indicate distinct behavioral patterns for some of the limpets in Port Valdez. Tectura persona exhibits negative phototactic and strongly reclusive be- havior and feeds primarily after sunset and before sunrise. During low tides in daylight hours, 7. persona typically aggregates under rocks or in crevices near boulder bases. Tectura fenestrata becomes abundant at tidal heights where 7. persona abundance decreases, and inhabits areas such as small crevices, the undersides of rocks in cobble beaches, and spaces within mussel clumps which afford it additional protection against thermal stress and desic- cation. Unlike 7. persona, other limpet species (Lottia species and 7. fenestrata) did not exhibit negative pho- totactic behavior nor were they as strongly reclusive, but all limpets appeared to avoid extreme weather conditions by moving into crevices or under rocks. Limpet abundance data revealed only broad trends in abundance related to tidal height. Lottia species and T. fenestrata were observed from the lower to high tidal heights with greatest abundance in the lower and mid- intertidal regions, whereas T. persona was more common in the high intertidal zone (Figure 3). The abundance data show that juveniles generally appeared in early to mid summer predominantly at lower tidal heights (Figure 3). Acmea mitra and T. scutum were observed during the study in extremely low abundance at the lower tidal heights. Seasonal abundance trends were only marginally apparent as there appeared to be an increase in abundance through the summer to late fall with lower abundance of all limpets in the winter; however, the trends are not con- sistent (Table 2 and Figure 3). Means and 95% confidence intervals for abundance data did not reveal patterns re- flective of site-to-site differences. Correlation analyses of mean monthly limpet abundance (ind. m~’ per tidal height) revealed a significant (p < 0.0083) positive cor- relation between Lottia species and juveniles, positive correlations of T. fenestrata abundance to Lottia species and to juveniles, and low negative correlations of T. per- sona to T. fenestrata and to juveniles (Table 3). Corre- Page 298 26 “%e. 7M, SB: Y=2.4910+1.6254*X 22 “BS. AB: Y=4.2035+1.3178*X 0.,, Ba: Y=2.8483+1.6254*X ox 18 8 = a $ to 14 7 q e 3 4 Flic’ ea tj «+10 Bo Ao $ o ice $ ,e = ee: H . algoree als 5 Oo 6 cme = o Se ae A Le ¥g 9 2 0 1 2 3 4 5 CLIE XR K LEK KI IOOO OO oO 6 \. The Veliger, Vol. 43, No. 4 ° ed fo) ited ° ape Jal 8 es oo eee 8 ka: ¥ Sf ae Haag 2 : o ‘27 M4 "a a: s & ie: 3 spore a A z fl 8 9 10 11 12 13 Estimated Age (Years) Figure 5 Regression lines for estimated age and shell length of Tectura persona for the four sites in Port Valdez, Alaska. lation analysis between monthly average abundance of limpets (ind. m~ for each site) and environmental vari- ables revealed significant positive correlations (p < 0.0031) of T. fenestrata abundance to air and water tem- peratures and a negative correlation to salinity; other spe- cies did not show significant correlations to the environ- mental variables (Table 4). Tectura persona demonstrates a seasonally distinct re- productive cycle. Developing gametes occurred through- out most of the year from spring to fall (Figure 4). Ripe gametes were observed at all sites in late August or Sep- tember, and spawning generally began in October. Lim- pets spawned at similar times throughout the fjord. Some spawning was observed in spring and summer by limpets larger than 11 mm. Regression analyses of growth data indicate significant relationships between age and length for 7. persona. Shell lengths varied from 2.7 mm to 22.9 mm with a mean of 11.2 mm, and estimated age ranged from 1 to 12 years with a mean of 5.3 years. Three separate regression lines were necessary to describe the age to length relationship for the four sites (Table 5, Figure 5). The regression equa- tions fit to 7M and SB length data were coincident. The intercept of the equation for the B4 was significantly higher (p < 0.05) than the equation for 7M and SB, in- dicating a slightly larger limpet population. The higher intercept and lower slope of the regression model for the AB site suggests larger juvenile limpets may be recruiting into the interidal population but that they are growing more slowly (in length) than at the other three sites. Re- gression analyses demonstrated that the relationships of both length and age to tidal height were not significant. DISCUSSION Inferences based on limpet abundance data agree with conclusions reached by other investigators. Field obser- vations suggest that T. persona is reclusive and demon- strates negative phototactic behavior, as also noted for this species by Lindberg et al. (1975). Also, the distri- bution of the six intertidal limpets within Port Valdez shows similarities to those reported for these species else- where along the Pacific coast (Fritchman, 1961:b, c, 1962; Lindberg, 1981, 1982). While all limpets are ex- posed to low salinities (approaching 0%c) in summer, and low temperatures in winter, only 7. fenestrata demon- strates a relationship between abundance values and en- vironmental parameters. Changes in abundance of T. fe- nestrata with temperature suggests increased cryptic be- havior and/or migration down the shore to avoid exposure to subfreezing air temperatures during low tides in winter. Tectura fenestrata and T. persona in San Francisco Bay, California, USA, are reproductively active (includes developing stages as well as spawning activity) during A. Blanchard & H. M. Feder, 2000 winter but are quiescent in summer when water temper- atures are high (Fritchman, 1961b, 1962). In Port Valdez, near the northern extreme of the range of T. persona, this limpet is reproductively active year-round with gonad de- velopment in spring and summer and spawning primarily from November through January (including the period of coldest temperatures; Figure 2). Large limpets (> 11 mm) spawn in summer as well, which indicates that they retain sufficient energy stores following winter to fuel gamete maturation and spawning in summer. It is presumed that smaller limpets do not retain the necessary energy stores for spawning in summer, and feeding in summer is re- quired to provide sufficient energy for the final stages of gametogenesis prior to spawning in winter. Assessment of fresh tissues from T. fenestrata also indicated year- round reproductive activity with fall and winter spawning (Feder et al., 1992). The summer water temperatures in Port Valdez are considerably lower than the summer wa- ter temperatures of California, and infrequently approach the critical water temperature (approximately 13°C) con- sidered to limit reproduction of 7. fenestrata and T. per- sona by Fritchman (1962). This year-round reproductive period for these two limpets in Port Valdez lends support to the conclusions of Fritchman that high water temper- atures (> 13°C) limit their reproductive activity (See also Underwood, 1979; Branch, 1981). In the southern range of these limpets, reduction of food resources in hot sum- mer months (e.g., desiccation of unicellular algae grow- ing as a film on bare surfaces; Cubit, 1984) may be im- portant in limiting reproduction but such conditions are rare in Port Valdez. In contrast, reproductive processes are strongest in winter, in spite of the lack of food re- sources, suggesting that food limitations have minimal influences on the reproductive cycle of T. persona and T. fenestrata. Similar to T. persona and T. fenestrata, year-round re- productive activity (including gametogenic development and spawning) is also described for the mussel Mytilus trossulus (Gould, 1850) in Port Valdez (Blanchard & Fed- er, 1997). Clarke (1987) noted that in polar regions, growth and reproduction are limited by the food supply, and gonad production may require two summer periods for completion. Likewise, it is possible that in addition to the lack of temperature constraints, the extended devel- opmental periods observed in T. persona, T. fenestrata, and M. trossulus may reflect responses to the climatic patterns in Port Valdez that are similar to but less extreme than adaptations described for polar organisms. Regression of shell growth against estimated age in- dicates a slow-growing T. persona population with mod- erate longevity. While the shell length to estimated age relationship of this species from Port Valdez appears to exhibit a slight curvature reflective of a nonlinear growth pattern, no concave models adequately fit the data. Thus it is concluded that the growth relationship of adult 7. persona in Port Valdez is nearly linear over the range of Page 299 lengths recorded. The moderate slopes (e.g., growth rate coefficients; Table 5) of the estimated regression func- tions of 1.3 to 1.6 and an observed maximum age of 12 years agrees in principle with Branch (1981) who indi- cated that limpets (including T. [Notoacmea] persona and other lottid limpets) with low growth coefficients live lon- ger. Studies of limpet shell growth indicate it is not un- common for differences in shell growth rate and shell size to occur, due to responses by limpets to natural environ- mental conditions (Branch, 1981). The differences in shell growth rate and shell sizes observed between sites in this study are well within the range of differences ex- pected as a response to natural conditions (e.g., Branch, 1981; Hobday, 1995). Determination of annual growth rings appears to be a valid aging tool for limpets in Prince William Sound, although marking studies were not con- clusive in demonstrating errors in estimated ages. How- ever, Kenny (1968) concluded that the dominant shell ridges of 7. persona from Oregon, USA, were annual growth rings. Additionally, shell growth ring analysis was successfully used for aging the barnacle Semibalanus bal- anoides (Linnaeus, 1767) in Port Valdez (Rucker, 1983), a number of bivalve species in Prince William Sound, including the mussel M. trossulus (Feder & Keiser, 1980; Blanchard & Feder, 2000), the littleneck clam Protothaca staminea (Conrad, 1857) (Paul & Feder, 1973), and the soft-shell clam Mya arenaria Linnaeus, 1758 (Feder & Paul, 1974), as well as eight other bivalve species in the southeastern Bering Sea (McDonald et al., 1981). In all of the latter Alaskan species, growth checks on shells reflected cessation of growth from reduced food resources associated with harsh winter conditions. There is no obvious relationship between shell length or estimated age and tidal height for 7. persona in Port Valdez. In a study of another limpet of the family Lotti- dae, L. digitalis, Hobday (1995) found few small limpets in a sheltered area along the California coast, compared to a more exposed area nearby. That study suggested that migration of larger limpets across the exposure gradient was the main source of recruitment to the sheltered re- gion. There may have been decreased survival for juve- nile and small limpets in the sheltered area as a result of reduced wave splash. In Port Valdez, small and large adult 7. persona were distributed throughout the mid to high tidal ranges and there was no evidence of reduced survival of smaller adults in the higher tidal regions. Compared to the study site in California (Hobday, 1995), the distribution of limpets in Port Valdez may be a result of reduced risk of desiccation for the smaller individuals due to local environmental conditions (e.g., shading of sites, cloudy weather, and a wet climate) although inter- specific habitat tolerances may explain some of the dif- ferences between the two studies. However, juveniles ap- pear in spring and early summer in Port Valdez and there did appear to be a movement of juveniles (< 5 mm) up- Page 300 shore with increased size as juveniles were much less common at higher tidal levels. One objective of the present study was to compare the distribution of limpets at a site within the marine terminal area (Station B4) with the distribution of limpets at sites outside the terminal area. After the oil spill by the Exxon Valdez in 1989, Highsmith et al. (1996) and Hooten & Highsmith (1996) observed decreased numbers of limpets on contaminated beaches. Additionally, Liu & Morton (1998) determined that limpets (Patelloida species) in highly polluted waters (primarily untreated sewage) al- locate more energy to reproduction, occur in lower den- sities, and have larger shell sizes than limpets in unaf- fected regions. In the present study, even when the B4 site was sampled 1 month following a minor crude oil spill in the terminal area (January 3, 1989, Alyeska Pipe- line Service Company, personal communication), limpet abundance values at B4 were similar to those of the other sites. In general, the abundance and reproductive patterns of limpets at B4 were no different from these parameters at the 7M, AB, and SB sites. The slightly increased shell lengths of T. persona at B4 are well within the range of differences expected as a response to natural environ- mental conditions. Other intertidal investigations in the vicinity of B4 demonstrated a relatively robust commu- nity at B4 (Feder & Bryson-Schwafel, 1988; Jewett et al., 1993). Acknowledgments. This research was initiated by Carol Klumpp prior to her death in 1990. We thank Kris McCumby for con- tinuing the field and laboratory portions of the study. Dr. Robert Benda assisted with fieldwork. We appreciate the help of Dr. David Lindberg with limpet identifications. We thank Dr. Lind- berg and an anonymous reviewer for their helpful and enlight- ening comments. We are grateful to the additional technicians and students who assisted with this study. Funding was provided by Alyeska Pipeline Service Company. LITERATURE CITED BLANCHARD, A. & H. M. FEpDgER. 1997. Reproductive timing and nutritional storage cycles of Mytilus trossulus Gould, 1850, in Port Valdez, Alaska, site of a marine oil terminal. The Veliger 40(2):121—130. BLANCHARD, A. & H. M. Feber. 2000. Shell growth of Mytilus trossulus Gould, 1850, in Port Valdez, Alaska. The Veliger 43(1):34—42. BRANCH, G. M. 1981. The biology of limpets: physical factors, energy flow, and ecological interactions. Oceanography and Marine Biology: an Annual Review 19:235—380. CLARKE, A. 1987. Temperature, latitude and reproductive effort. Marine Ecology Progress Series 38:89—99. Cusit, J. D. 1984. Herbivory and the seasonal abundance of algae on a high intertidal rocky shore. Ecology 65(6):1904— 1917. Feber, H. M. & A. J. PAUL. 1974. Age, growth and size-weight relationships of the soft-shell clam, Mya arenaria, in Prince William Sound. Proceedings of the National Shellfish As- sociation 64:45—52. FEDER, H. M. & G. E. KEISER. 1980. Intertidal biology. Pp. 143— 224 in J. M. Colonell & H. K. Stockholm (eds.), Port Valdez, The Veliger, Vol. 43, No. 4 Alaska. Environmental Studies, 1976-1979. Institute of Ma- rine Science Occasional Publication No. 5, University of Alaska: Fairbanks, Alaska. Feper, H. M. & B. BRYSON-SCHWAFEL. 1988. The intertidal zone. Pp. 117-164 in D. G. Shaw & M. J. Hameedi (eds.), Envi- ronmental Studies in Port Valdez, Alaska. Springer-Verlag: New York. Feber, H. M., K. McCumsy & S. C. Jewett. 1992. A long-term intertidal monitoring program for Port Valdez, Alaska 1990— 1991. Component 3—Biological study of the dominant in- tertidal limpet Tectura persona and the distribution and abundance of other intertidal limpets in Port Valdez, Alaska. Final Report to Alyeska Pipeline Service Company. Uni- versity of Alaska Fairbanks. 298 pp. FRANK, P. W. 1965a. Growth of three species of Acmaea. The Veliger 7(4):201—202. FRANK, P. W. 1965b. Biodemography of an intertidal snail pop- ulation. Ecology 46(6):83 1-844. FRITCHMAN, H. K. 1961la. A study of the reproductive cycle in the California Acmaeidae (Gastropoda). Part I. The Veliger 3(3):57-63. FRITCHMAN, H. K. 1961b. A study of the reproductive cycle in the California Acmaeidae (Gastropoda). Part II. The Veliger 3(4):95-101. FRITCHMAN, H. K. 1961c. A study of the reproductive cycle in the California Acmaeidae (Gastropoda). Part III. The Veliger 4(1):41-47. FRITCHMAN, H. K. 1962. A study of the reproductive cycle in the California Acmaeidae (Gastropoda). Part IV. The Veliger 4(3):134-140. GOERING, J. J., W. E. SHtets & C. J. PATTon. 1973. Nutrient cycles. Pp. 223-250 in D. W. Hood, W. E. Shiels & E. J. Kelley (eds.), Environmental Studies of Port Valdez. Insti- tute of Marine Science Occasional Publication No. 3, Uni- versity of Alaska: Fairbanks, Alaska. HIGHsMITH, R. C., T. L. RUCKER, M. S. STEKOLL, S. M. SAUPE, M. R. LINDBERG, R. N. JENNE & W. P. ERIKSON. 1996. Impact of the Exxon Valdez oil spill on intertidal biota. Pp. 212— 237 in S. D. Rice, R. B. Spies, D. A. Wolfe & B. A. Wright (eds.), Proceedings of the Exxon Valdez Oil Spill Sympo- sium. American Fisheries Society Symposium 18. Hospay, A. 1995. Body-size variation exhibited by an intertidal limpet: influence of wave exposure, tidal height and migra- tory behavior. Journal of Experimental Marine Biology and Ecology 189:29—45. Hoop, D. W., W. E. SHIELS & E. J. KELLEY. 1973. Environmental studies of Port Valdez. Institute of Marine Science Occa- sional Publication No. 3, University of Alaska: Fairbanks, Alaska. Hooten, A. J. & R. C. HIGHSMITH. 1996. Impacts on selected intertidal invertebrates in Herring Bay, Prince William Sound, after the Exxon Valdez oil spill. Pp. 249-270 in S. D. Rice, R. B. Spies, D. A. Wolfe & B. A. Wright (eds.), Proceedings of the Exxon Valdez Oil Spill Symposium. American Fisheries Society Symposium 18. JEWETT, S. C. & H. M. FEDER. 1977. Biology of the harpacticoid copepod Harpacticus uniremis Kréyer on Dayville Flats, Port Valdez, Alaska. Ophelia 16:111—129. Jewett, S. C., H. M. FEDER & A. BLANCHARD. 1993. A long- term intertidal monitoring program for Port Valdez, Alaska 1990-1992. Final report to Alyeska Pipeline Service Com- pany. University of Alaska Fairbanks. 275 pp. Kenny, R. 1968. Growth characteristics of Acmaea persona Es- chscholtz. The Veliger 11(4):336—339. A. Blanchard & H. M. Feder, 2000 LINDBERG, D. R. 1981. Acmaeidae. Boxwood Press: Pacific Grove, California. 122 pp. LINDBERG, D. R. 1982. Taxonomic notes on members of the ge- nus Colisella from the North Pacific Ocean including a de- scription of a new species from Alaska (Gastropoda: Ac- maeidae). The Wassman Journal of Biology 40(1—2):48—58. LINDBERG, D. R. 1986. Name changes in the ‘“‘Acmaeidae.”’ The Veliger 29(2):142-148. LINDBERG., D. R., M. G. KELLOG & W. E. HuGuHEs. 1975. Evi- dence of light reception through the shell of Notoacmea per- sona (Rathke, 1833) (Archaeogastropoda: Acmaeidae). The Veliger 17:383—386. Liu, J. H. & B. Morton. 1998. The impacts of pollution on the growth, reproduction and population structure of Hong Kong limpets. Marine Pollution Bulletin 36(2):152—158. Lowe, D. M., M. N. Moore & B. L. BAyne. 1982. Aspects of gametogenesis in the marine mussel Mytilus edulis L. Jour- nal of the Marine Biological Association of the United King- dom 62:133-145. McDONALD, J., H. M. FEDER & M. HoBerG. 1981. Bivalve mol- lusks of the southeastern Bering Sea. Pp. 1155-1204 in D. W. Hood & J. Calder (eds.), The Eastern Bering Sea Shelf: Page 301 Oceanography and Resources, Volume II, U. S. Department of Commerce. NETER, J., W. WASSERMAN & M. H. KuTNer. 1990. Applied Lin- ear Statistical Models. Irwin: Homewood, Illinois. 1181 pp. Orton, J. H., A. J. SourHWARD & J. M. Dopp. 1956. Studies on the biology of limpets. II]. The breeding of Patella vulgata L. in Britain. Journal of the Marine Biological Association of the United Kingdom 35:149-176. PauL, A. J. & H. M. Feber. 1973. Growth, recruitment and dis- tribution of the littleneck clam, Protothaca staminea, in Ga- lena Bay, Prince William Sound, Alaska. Fisheries Bulletin 71:665-677. RucKER, T. L. 1983. The life history of the intertidal barnacle Balanus balanoides (L.) in Port Valdez, Alaska. M.S. Thesis, University of Alaska Fairbanks. 251 pp. Situ, J. E. (ed.). 1968. Torrey Canyon Pollution and Marine Life. Cambridge University Press: New York. 196 pp. THompson, J. B. 1980. Distribution and population dynamics of the limpet Patella vulgata L. in Bantry Bay. Journal of Ex- perimental Marine Biology and Ecology 51:145-171. UNDERWOOD, A. J. 1979. The ecology of intertidal gastropods. Pp. 111-210 in EF S. Russell & C. M. Yonge (eds.), Ad- vances in Marine Biology. Academic Press: New York. The Veliger 43(4):302—312 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 Development and Anatomy of Nitidiscala tincta (Carpenter, 1865) (Gastropoda: Epitoniidae) R. COLLIN Committee on Evolutionary Biology, University of Chicago, Culver Hall, 1025 E. 57th Street, Chicago, Illinois 60637, USA and Department of Zoology, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605, USA (e-mail: rcollin@ midway.uchicago.edu; phone: (312) 922-9410 x 578) Abstract. The epitoniids (Gastropoda: Ptenoglossa) are one group for which there are no current well-supported hypotheses of relationships. Herein I describe the development and adult anatomy of an epitoniid, Nitidiscala tincta, and discuss intra-familial variation in characters used for higher level systematics. Female N. tincta lay 75 wpm eggs in clusters in sand-covered capsules connected together along a mucus strand. The first two cleavages are synchronous, appear equal, and produce a polar lobe. Gastrulation is by epiboly and produces a trochophorelike stage with embryonic kidneys. At hatching the larvae are 125 wm long and have a small velum, heart, eyes, and a well-developed, black pigmented mantle organ. The shell is right-handed and slightly hydrophobic. The mantle cavity contains a gill with triangular lamellae, purple hypobranchial gland, and a triple-ridged osphradium. The acrembolic probosis ends in a robust pair or jaws slightly anterior to the split odontophore, a ptenoglossan radula, and two pairs of salivary glands. The presence of both caenogastropod and heterobranch developmental synapomorphies suggests that ideas about the evolution of development among gastropods may need to be reevaluated. INTRODUCTION Recent interest in high level gastropod systematics has focused on the relationships among well-known, purport- edly monophyletic groups, using either exemplar species or composites of several disparate species (Haszprunar, 1988; Mikkelsen, 1996; Ponder & Lindberg, 1997). Some poorly known groups like deep-sea limpets and lower het- erobranchs have also received considerable attention due to their perceived phylogenetic importance and the avail- ability of recent detailed anatomical studies (Haszprunar, 1988; Healy, 1993; Ponder & Lindberg, 1997). However, there are still many groups whose relationships are not well understood. The Epitoniidae is one such group. What little is known about the biology of epitoniids has been used to group them with either heterobranchs (Gosliner, 1981; Habe & Kosuge, 1966; Robertson, 1985) or caen- ogastropods (Fretter & Graham, 1994; Haszprunar, 1985). Robertson (1983a b, 1985) used reproductive and de- velopmental data to argue for affinities of epitoniids and groups now considered to be lower heterobranchs. He pointed out that Epitonium Roding, 1798, species possess two putative heterobranch synapomorphies (Robertson, 1985): they lay eggs that are connected by chalazae (Rob- ertson, 1973; but see Robertson, 1985 for a different opinion), and their larvae have pigmented mantle organs (PMO). Robertson (1983b) also reported that the larvae of E. albidum (d’ Orbigny, 1842) have hydrophobic larval shells, another characteristic of heterobranchs (Collin, 1997). Current opinion favors a close relationship between ep- itoniids and caenogastropods based on anatomical and ul- trastructural similarities (Haszprunar, 1985; Healy, 1994; Ponder & Lindberg, 1997). Published discussions of high- level gastropod systematics group epitoniids with janthin- ids in the Janthinoidea (synonym: Epitonioidea; see Pon- der & Warén, 1988), because members of these families share a ptenoglossan radula, an accessory pair of salivary glands, feed on cnidarians, and produce a purple secretion when disturbed (Fretter & Graham, 1994). The proto- conchs are also strikingly similar. The Janthinoidea are grouped with the Triphoridae, Triforidae, Cerithiopsidae, Aclidae, and Eulimidae, as the Ptenoglossa on the basis of characters listed in Table 1. In recent phylogenetic analyses of the Gastropoda, a composite triphorid was used to represent the Ptenoglossa (Ponder & Lindberg, 1996, 1997). However, Ponder & Lindberg (1997) stated that the Ptenoglossa is ‘‘an almost certainly paraphyletic or polyphyletic taxon.’’ This hypothesis cannot yet be tested because complete anatomical studies of most pten- oglossan groups are not currently available. The characters listed in Table 1 are not optimal for R. Collin, 2000 Table 1 Taxonomic distribution of characters traditionally used to support the “‘ptenoglossa.”’ Characters* Epitoniidae Janthinidae Aphalic males yes/no yes Spermatozeugmata yes yes Protandrous hermaphrodites yes yes Ptenoglossiate radula yes yes Divided radula/odontophore yes yes Stylets yes yes Medial tooth yes/no no Adult purple gland yes yes Larval PMO yes yes Acrembolic proboscis yes no Cuticularized esophagus yes yes Zygoneury yes/no yes Osphradium ridges 1 or 3 1 or 2 Open pallial oviduct yes no Pairs of salivary glands 2 2) Esophageal glands no no ““Beaked”’ larval shell no no Statocysts yes no Page 303 Eulimidae Cerithiopsidae Triphoridae no yes yes no yes yes some no no yes taenoglossan sometimes ? no no no no no no yes yes no no no no no no yes yes yes yes yes no yes ? no 3 or | ? I yes/no yes yes l 1 1 no? maybe maybe no yes yes yes u 2 * Characters are coded based primarily on the following papers. Epitoniids: Bouvier 1886; Healy 1994; Taki 1956, 1957; Thiele 1928. Janthinids: Bouvier 1886; Graham 1965; Healy 1994; Laursen 1953; Thiele 1928; Wilson & Wilson 1956. Eulimids: Warén 1983a, b. Triphorids: Fretter 1951; Healy 1990; Kosuge 1966; Marshall 1983. Cerithiopsids: Fretter 1951; Healy 1990. phylogenetic analysis for several reasons. First, the vari- ation within each group has not been investigated and may not be represented adequately. For example, most accounts of epitoniid radulae state that the central tooth is absent. Examination of Couthouyella Bartsch, 1909, however, shows that this is not the case in all epitoniids (Warén, 1980). Second, because so little is known about any one species, each group is coded as a composite of characters from many species that may not be closely related. Finally, homology assessments may based on very littke comparative information. For example, both epitoniids and janthinids clearly have two pairs of sali- vary glands, whereas most caenogastropods have only one. However, cerithiopsids have been described as pos- sessing a single pair (Fretter, 1951) in which one gland connects to the anterior esophagus and the other connects to the posterior esophagus. The morphology and _ultra- structure of the two glands are different. This condition is interpreted as being a single pair in which the two glands have differentiated from each other (Fretter, 1951), but it could also represent a condition in which there were two pairs of glands with subsequent loss of one gland in each pair. Interpretation of osphradial morphology illus- trates a similar situation: epitoniids and janthinids have either a single simple ridge or multiple parallel ridges. Janthina Réding, 1798, species have either one or two ridges, and epitoniids are reported to have one or three ridges. Triphorids, however, have a single thick ridge with a medial groove (Kosuge, 1966). It is unclear if this con- dition should be interpreted as a different character state Or an intermediate form. Examination of more species will significantly increase our ability to refine these char- acters and to make well-informed homology assessments. Herein I examine the development and adult morphol- ogy of Nitidiscala tincta (Carpenter, 1865). This study was undertaken (1) to provide the first description of early development for a member of this family, and (2) to pro- vide descriptive morphological data that can be used as a basis for comparative and phylogenetic studies of epi- toniids or Ptenoglossa. MATERIALS AnD METHODS Adult Nitidiscala tincta were collected from the mid- intertidal zone near Santa Barbara, California (from Al- egria [34°28’N, 120°17’W] and Coal Oil Point [34°05'N, 120°10’W)) in July 1997. They were collected by hand during low tide from sand around the base of the anem- one Anthopleura elegantissima (Brandt 1835). Species were identified using DuShane (1979), and vouchers are deposited at the Field Museum of Natural History in Chi- cago (FMNH 282448). Animals were kept at ambient sea temperature (16—18°C) in custard dishes, with a layer of sand and one or several host anemones. Feeding behavior was observed under a dissecting microscope. The water was changed and the eggs were collected daily. Egg strings were kept in glass custard dishes or large petri dishes until the embryos hatched. One 11 mm-long fe- male was kept alone to determine if females can store sperm. Embryos were collected by pulling apart the capsules with forceps. Developmental stages were observed and Page 304 Figure 1 Adult N. tincta. Scale bar = 4 mm. photographed every hour during cleavage and every day thereafter, at a total magnification of 100—200 with an Olympus compound microscope. Upon hatching, the lar- vae were measured and transferred to custard dishes with a density of about 1—2/10 ml. They were fed the unicel- lular brown alga Jsochrysis galbana, and the water was changed every other day. Larvae were maintained this way for 2 weeks, but no further attempt was made to raise them to settlement. Standard Prussian blue staining (Clarke, 1973) was used to detect embryonic albumin uptake. Embryos were carefully excapsulated and incubated in a solution of fer- ritin in seawater for 1—2 hours. They were then relaxed, fixed in 10% formalin, rinsed, and stained with HCI and potassium ferrocyanide solutions. In the presence of iron (from the ferritin) a blue product is formed. Staining was clearly visible with whole mount light microscopy. A negative control, for which the embryo was placed into a solution of seawater instead of ferritin was used in all staining experiments. Adult animals were fixed in formalin and decalcified prior to staining with toluidine blue and dissection under a Wild M4 dissecting microscope. Tissues for SEM ob- servations were dissected from fixed, decalcified animals, dehydrated in ethanol followed by Hexamethyldisalizine, and viewed with an Almray scanning electron micro- scope. RESULTS Adult N. tincta (Figure 1) are abundant on large patches of Anthopleura elegantissima in the mid-intertidal zone (Breyer, 1982). They occur amongst closely packed anemones buried in the sand, or occasionally stuck to an anemone’s column wall by spirocysts. In the laboratory, snails usually remained buried completely in the sand un- less they were feeding. Adults were attached to the sub- strate, their egg strings, and each other by a sticky mucus The Veliger, Vol. 43, No. 4 Figure 2 Egg capsules of N. tincta. Scale bar = 1 mm. attachment thread secreted from the posterior sole of the foot. Attachment threads of several individuals were often entangled. Egg strings were commonly attached to anem- ones in the field, and egg strings from several females were usually deposited in a single tangle. Anemones do not respond when they contact the egg strings with their tentacles or oral disk. Development Female N. tincta lay egg strings which consist of mul- tiple sand agglutinated capsules (Figure 2) arranged along a mucus thread, which is similar to and often connected to the attachment thread. Each capsule consists of a tight cluster of eggs surrounded by a capsule of tightly packed sand grains held together by a thin layer of mucus. The mucus thread which connects the capsules passes through the middle of each capsule, and the sand, eggs, and mu- cus can be removed without damaging the thread. In the laboratory, the isolated female continued to lay fertilized eggs for at least 16 days without reduction in mean eggs/ capsule or egg viability. On average this female laid one egg string with 149 capsules every 2 days. There were 10—60 eggs per capsule (mean = 32.9, SD = 13.1, n = 80 capsules, five each from 16 egg strings from different mothers). The number of eggs per capsule did not vary within an egg string, but did vary signifi- cantly among strings from different females (one-way ANOVA: F = 50.3, df = 15, p < 0.001). The mean outer diameter of 32 capsules from three females was 1.02 mm (SD = 0.157) and also varied among strings from differ- ent females (one-way ANOVA: F = 52.2, df = 2, p < 0.001). The small 75.3 wm (n = 62 from five females, SD = 1.7) white eggs were deposited while the germinal vesicle was still visible as a clear area in the otherwise opaque cytoplasm. Egg size did not vary among capsules, egg strings, or females. All eggs within a capsule devel- R. Collin, 2000 Table 2 Developmental time table for WNitidiscala tincta at 16-18°C Age Stage 7 hours First cleavage. 9 hours 2-cells. 10—11 hours 4-cells. 12 hours 3rd cleavage. 24 hours Late cleavage. 2-3 days Gastrulation. 3.5—4 days Gastrulation complete, mouth visible, some slight ciliation. 4.5 days ‘“‘Trochophore”’ stage: Velum anlagen visible, larval kidneys. 5.5 days Some shell growth, foot and operculum visi- ble. 7 days Early veliger morphology: Shell sculpture, long velar cilia, black PMO, and statocysts visible. 9 days Dark color on shell sutures appears. 11 days Heart beats slowly; eyes and right tentacle are visible. 12 days Hatching at 125 pm. oped synchronously, but the developmental stages were slightly staggered along an egg string. After the germinal vesicle breaks down, the eggs have a uniformly granular appearance and two polar bodies are extruded. A developmental schedule of the following events is given in Table 2. The first two cleavages are both synchronous, appear to be equal, and produce a small polar lobe (Figure 3A). The third cleavage is dif- ficult to see and produces four clear micromeres that are close to the size of the macromeres (Figure 3B). Subse- quent cleavages are not quite synchronous. Although the membrane between the macromeres and micromeres is difficult to see, one of the macromeres rounds up and compacts slightly out of phase with the other macro- meres. This asynchrony may begin as early as the third cleavage. Subsequent cleavage produces a round blastula that gastrulates by epiboly (Figure 3C): The clear micro- meres slowly grow around the more opaque yolky mac- romeres. There is no invagination. After gastrulation, the embryo is slightly elongate, with a mouth about one-third of the way behind the apical tuft. A small ridge, the head- foot anlage, forms anterior to the mouth, and two small, clear embryonic kidneys appear lateral to the mouth (Fig- ures 3D, E). The kidneys are equal in size and both take up ferritin. At this stage the embryo is slightly ciliated. The head-foot anlage gradually differentiates into a velar ridge and the foot, while the shell and operculum develop. After 8 days the statocysts are well developed, and the black PMO become visible on the dorsal right side, just posterior to the velum. Two areas of reddish pigment also develop on the shell, along the suture. At no stage is there Page 305 a head vesicle. After 11 days the heart beats weakly, both eye spots are present, and the right tentacle is distinct. However the viscera is still an indistinct yolky mass. At this stage, Prussian blue staining shows that the left em- bryonic kidney has become much smaller and is difficult to detect in whole mounts, but still takes up ferritin. The only structure to stain on the right side is the PMO, which stains strongly in both the ferritin treatment and the neg- ative control. The strong staining of the PMO might ob- scure the right embryonic kidney, if it is still functional. Planktonic feeding larvae hatch after 13 days with a shell length of 124.7 wm (n = 89 from seven females, SD = 3.3; Breyer’s [1982] statement that they hatch at 72 wm is in error) (Figure 3F). The shell is right handed. The velum and foot are small and pigmentless and the viscera are still not clearly differentiated, although there is a clear muscle attachment at the shell apex. The eyes and heart are well developed, but the tentacles are small. The velum does not project much from the body wall and is made up of few large cells, similar in appearance to those of vetigastropod veligers. The larval shells are hy- drophobic and occasionally become trapped in the surface tension, although larvae generally remain near the bottom of the culture dish. Nitidiscala tincta veligers have been reported to live in culture for at least 2 months without settling (Smith & Breyer, 1983). Because animals were not raised through metamorphosis it was not possible to determine if the larval PMO is homologous to the adult hypobranchial gland as discussed by Robertson (1983b, 1985). Feeding Snails of all sizes (S—14 mm) were observed feeding on anemones in the laboratory. The proboscis is everted slowly until the expanded distal end is fully extended (Figure 4). The fully everted proboscis is approximately 1.5 times the length of the shell. The jaws, which are visible through the transparent wall of the proboscis, lie in the distal expansion of the extended proboscis. The extended proboscis waves around and probes the anem- one. When the mouth contacts the end of a tentacle, it slips around the tentacle and works its way toward the base of the tentacle, engulfing it (Figure 4). There is a faint pumping motion on the ventral side of the proboscis just posterior to the distal expansion, and the engulfed tentacle appears to shrink a little. Then with a sudden smooth motion the distal end of the proboscis contracts laterally and severs the tentacle along the line of the tooth plates. The severed tentacle can be seen through the pro- boscis wall, moving toward the snail’s head, and the pro- boscis quickly retracts. The anemone shows a clean di- agonal incision on the tentacle stub where the tentacle was removed. Although snails usually engulf tentacles from the end, they sometimes surround them in the mid- dle, folding them in half as the proboscis covers them, The Veliger, Vol. 43, No. 4 Figure 3 Developmental stages of N. tincta. A. Two-cell stage showing the polar bodies at the animal pole and the polar lobe at the vegetal pole. B. Later cleavage showing that the micromeres and macromeres are similar in size. C. Gastrula, after cell compaction. D. Early ‘‘trochophore”’ stage, showing the mouth and the embryonic kidneys. E. Late ‘“‘trochophore”’ stage showing embryonic kidneys. FE Hatchling larva showing the distinct black PMO. Key: k = embryonic kidneys, pb = polar bodies, pl = polar lobe. Scale bar = 40 wm. and sometimes they attack the tubercles on the anemone’s column. In all cases that I observed, the snails attached the proboscis to some projection of the anemone’s body wall. At no time did I observe the snails tearing bits from the anemone, engulfing them whole, or inserting the pro- boscis into the a hole in the anemone’s body wall as has been reported in other descriptions of epitoniid feeding (den Hartog, 1987; Perron, 1978; Robertson, 1963, 1983b; Salo, 1977; Thorson, 1957). Such differences in feeding biology may be associated with the anatomical variation discussed below. Morphology External Morphology. The N. tincta collected for this study ranged in shell length from 5—14 mm. The shells are white with a distinctive purple marking overlying the hypobranchial gland along the top of the body whorl. The operculum is brown, uncalcified, ovate, and coiled. The visible body is white, with a rectangular foot, short ta- pered tentacles with an eye positioned dorso-laterally at the base of each. The snails are often attached to egg strings, anemones, or the substrate by a sticky string that originates in a medial groove that runs along the ventral side of the foot. The proboscis opening lies between and slightly below the tentacles. The viscera extend about four or five whorls into the shell. When removed from the shell, the dark coloration of the hypobranchial gland, the gills, and the osphradium can be seen clearly through the mantle. Mantle Cavity. The mantle cavity is similar to those of other coiled caenogastropods and extends one whorl back from the aperture. The gill lamellae are triangular and the gill extends posteriorly to the end of the mantle cavity (Figure 5). Dorsal and parallel to the gill is the hypob- ranchial gland, a long ridge of large cells full of a purple exudate, which extends back about 75-85% of the first R. Collin, 2000 Figure 4 Adult N. tincta feeding on anemone. A. The proboscis probes the anemone, and B. slips around a tentacle. C. The proboscis contracts to cut the tentacle. Key: sh = shell, pb = proboscis. whorl. In fixed material the gland becomes very dark, almost black, and the pigment takes on a crystalline ap- pearance. The pigment stains the edge of the gills next to the hypobranchial gland, and the mantle anterior to the hypobranchial gland is speckled brown. The osphradium is composed of three simple ridges that run along the ventral edge of the gill and curve ventrally at their an- terior end. The two external ridges have evenly spaced transverse striations and connect to each other around the anterior edge of the middle ridge (Figure 5). The central ridge is smooth. The distal portion of the intestine sits Page 307 Figure 5 Diagram of the reflected mantle cavity. Key: g = gill, hg = hypobranchial gland, i = intestine, os = osphradium, po = pallial oviduct. between the pallial oviduct and the hypobranchial gland but extends only halfway to the aperture (Figure 5). The columellar muscle also extends the length of the first whorl. The heart and kidney lie just posterior to the man- tle cavity. The heart is typically monotocardian and is surrounded by the flat round kidney. Alimentary System. The most prominent feature of the alimentary system is the acrembolic proboscis. The pro- boscis is surrounded by a layer of circular muscles which serves as an attachment for a complex assortment of re- tractor and extensor muscles. In the fully extended posi- tion, the jaws lie at the distal tip of the proboscis with the radula immediately behind them. In the fully retracted position, the jaws are at least a third of the way back from the opening into the proboscis introvert (Figure 6). The two large triangular jaw plates are surrounded by a muscular sheath (Figure 7A, B). Each jaw is made of many individual lamellae that can be seen easily with light microscopy to extend across the width of the jaw. The distal edge of each lamella projects through the sur- rounding tissue producing a row of fine serrations along Page 308 st cc Figure 6 Diagram of the proboscis and salivary glands with the overlying tissue and the complex assortment of retractor muscles removed. These structures are reflected to the animal’s left so that a lateral view of the proboscis is presented. The nerve ring is also dis- sected from the connective tissue in the head, but the ganglia are portrayed in situ. Key: bg = buccal ganglion, bgc = buccal gan- glia connectives, cc = cardiac caecum, j = jaws, Icg = left ce- rebral ganglion, lpg = left pleural ganglion, pdg = pedal ganglia, The Veliger, Vol. 43, No. 4 the edge of the jaw (Figure 7C). Immediately anterior to the jaws, embedded in the muscular wall of the oral tube are two small muscular stylet bulbs (Figure 6). Two fine cylindrical stylets project from the bulbs into the lumen of the proboscis. Ducts embedded in the wall of the esophagus run posterior from these bulbs to the salivary glands. Immediately posterior to the jaws is the divided odon- tophore and radula. The radula is as wide as it is long, has no central rachidian tooth, and is only slightly curved in the dorsal-ventral plane. The teeth vary in morphology; some have a hook on the distal end, whereas others end in a point (Figure 7D). There are a variable number of serrations along the edge proximal to the hook. The me- dial teeth are not as densely packed and have a more needlelike appearance, whereas the lateral ones are dense- ly packed and more clearly hooked (Figure 7D). There are two pairs of tubular salivary glands (Figure 6). The outer pair are relatively thick, slightly flattened, and have a rough appearance. Anteriorly, they attach to the buccal mass at the level of the buccal ganglia, pos- terior to the jaws and radula, but I was not able trace their ducts. The inner pair are long, smooth, thin coiled tubes that loop along the esophagus and attach near the buccal ganglia. Their ducts then run along the proboscis to the stylet bulbs. Posteriorly, all four glands are attached to the posterior proboscis in the same place. Posterior to the salivary glands, the posterior constric- tion of the esophagus connects to the cardiac pouch and the stomach. The stomach’s thin walls are covered inter- nally with numerous shallow folds. It is not cuticularized and there is no style sac. The intestine is simple and emp- ties into the back of the mantle cavity. Nervous System. The nervous system is similar to other typical streptoneurous gastropods. There are two distinct buccal ganglia at the base of the salivary glands (Figure 6). When the proboscis is retracted, they lie far back in the first whorl. The nerve ring lies in the animal’s head, directly behind the tentacles (Figure 6). All the ganglia, except the sub-esophageal ganglia lie on the dorsal left side of the esophagus, behind the left tentacle. The ce- rebral ganglia are large, round, and flattened, and the commissure is very short. There is a distinct nerve run- ning from each cerebral ganglion into the tentacle on the same side. The slightly elongated pleural ganglia lie be- hind and to the left of the cerebral ganglia and are tightly connected to them. The right pleural ganglion is oriented transversely, whereas the left one has an anterior-posterior orientation. The subesophageal ganglion lies against the &— rcg = right cerebral ganglion, rpg = right pleural ganglion, sb = stylet bulb, sbg = subesophageal ganglion, sg = salivary glands, sgd = salivary gland ducts, spg = supraesophageal gan- glion, st = stomach. R. Collin, 2000 WON Figure 7 SEM of the jaws and radula. A. View of the buccal cavity through the jaws that have been split open along the dorsal mus- cular connection. The natural opening is along the ventral edge (bottom in this view). The radula is at the base of the jaws. Scale bar = 100 pm. B. One jaw removed from the buccal mass. Ar- row indicates the serrations enlarged in C. Scale bar = 100 ym. C. Close-up view of the serrations where the underlying jaw extends from the tissue. Scale bar = 5 pm. D. The radula. The area with sparse teeth is more medial on the radula. Scale bar = 10 pm. Page 309 columella muscle under a thin layer of connective tissue on the far right side on the muscle. It does not connect to the right cerebral or pleural ganglia. The supraesopha- geal ganglion is connected to the right pleural ganglion by a short connective and lies to the left of and slightly below the other ganglia in the connective tissue just to the left of the columella muscle. The two oval pedal gan- glia are connected to the rest of the nerve ring by a large number of individual connectives that run from the pleu- ral and cerebral ganglia. They are deeply embedded in the foot musculature. Reproductive System. The gonad occupies the dorsal two-thirds of all the posterior whorls. In the female the pallial oviduct occupies the dorsalmost one-third of the body whorl. It is large, glandular, and slightly flattened. It is not differentiated into distinct regions and is open along its entire length on the dorsal side. Males are aphal- lic and produce large spermatozeugmata. DISCUSSION Development The development of N. tincta is unusual because it combines characters that were previously thought to be specific to caenogastropods with characters that were thought to be specific to heterobranchs (Table 3). Nitidis- cala tincta develops a distinct polar lobe and embryonic kidneys that are characteristic of caenogastropods. How- ever, the larva has a dark pigment spot that appears to be the same as a larval heterobranch PMO, and the larval shell is hydrophobic. None of these characters has been reported in non-heterobranch, non-caenogastropod snails, so it is unlikely that these characters are symplesiomor- phies. Epitoniid egg capsules are typically sediment- (sand or mud) covered mucus capsules that are strung together along a mucus thread (Robertson, 1983a b). However, Habea inazawai Kuroda, 1943, capsules have a reticulate surface and are not covered with sediment (Habe, 1943). There are no previous descriptions of early development to compare with my observations. The few accounts of epitoniid larvae describe them as small and having a prominent PMO (Habe, 1943; Thorson, 1946). All ob- served epitoniid larvae have right-handed shells, which brings into question the observation that Couthouyella is heterostrophic (Gosliner, 1981). Little is known about the embryology of other pten- oglossan gastropods (with the possible exception of some viviparous janthinids). Natarajan (1957) observed that hatching larvae of Janthina are small and have a distinct PMO but no eyes or tentacles. Cipriani et al. (1994) ob- served embryonic kidneys in cerithiopsid embryos. The larvae have a distinct dark spot in the viscera, which is not, however, in the appropriate position for a PMO (R. Cipriani, personal communication). The later stages of cerithiopsids and triphorids are, however, well known Page 310 The Veliger, Vol. 43, No. 4 Table 3 Distribution of developmental characters between caenogastropods and heterobranchs, and their condition in Nitidiscala tincta. Character Caenogastropod Heterobranch Cleavage polar lobe unequal Embryonic kidney yes no Larval PMO no yes Hydrophobic larval shell no yes Left handed larval shell no yes Large complex larvae* yes no N. tincta Reference polar lobe Freeman & Lundelius, 1992 van den Biggelaar & Haszprunar, 1996 yes Fioroni, 1966 yes Robertson, 1985 yes Collin, 1997 no Robertson, 1985 no Page, 1994 * Unlike the other characters, high larval complexity of caenogastropod larvae is a generalization and not a synapomorphy. (Lebour, 1933; Fretter & Pilkington, 1970). They have a large bilobed velum, complex shell sculpture, and a dis- tinct apertural notch. The protoconchs of janthinids, epi- toniids, and eulimids are all smooth, many (comprising three to four) whorls, and have a distinctive shape that is very different from the protoconchs of cerithiopsids and triphorids. Anatomy The anatomy of N. tincta as described herein also gen- erally agrees with the few previous descriptions of epi- toniid anatomy. These descriptions are difficult to com- pare because different techniques were used to examine different aspects of anatomy on different species. Studies that used basic dissections include the following: Taki (1956, 1957) described the morphology of the mantle cavity and alimentary systems of five Japanese genera (Epitonium, Amaea H. & A. Adams, 1853, Papyriscala de Boury, 1909, Gyroscala de Boury, 1887, and Acustis- cala de Boury, 1909); Bouvier (1886) focused on the ner- vous system of Scalaria communis Lamarck, 1822; and Warén (1980) and Gosliner (1981) commented briefly on Couthouyella striatula (Couthouy, 1839). Thiele (1928) made a more detailed study of Scala magellanica Philip- pi, 1845. Studies by Taki (1956, 1957) and Fretter & Graham (1994) focused mainly on the foregut anatomy. Like N. tincta, individuals of E. (Clathrus) clathrus Linnaeus, 1758, and the five Japanese genera have a long acrem- bolic proboscis with a poorly demarcated buccal mass, jaw, and two pairs of salivary glands. Taki’s studies show that there is, however, considerable variation in the rela- tive size of the jaws and radulae and the development of the salivary glands. In these previous studies the jaws are figured as being smaller or perhaps the same size as the radula. In N. tincta the jaws are clearly larger than the radula. Thiele (1928) described stylets connected to ducts that empty into the oral cavity. Fretter & Graham (1994) also observed these in E. clathrus, but Taki and Bouvier apparently did not observe these structures. I observed stylet bulbs in N. tincta which agree with Hochberg’s (1971) observation on the same species. These differenc- es may relate to differences in feeding behavior. Unfor- tunately morphological descriptions are not available for the species for which the diet and feeding habits have been described. The two prior reports of the epitoniid nervous system (Bouvier, 1886; Thielle, 1928) observed different degrees of zygoneury. In the species they studied, as in WN. tincta, the nerve ring lies anterior in the head directly behind the left tentacle, and the pedal ganglia are connected to the rest of the nerve ring by long connectives, as are the sub- and supraesophageal ganglia. Bouvier (1886) found the sub- and supraesophageal ganglia embedded in the body wall, which agrees with the location of both against the columellar muscle in N. tincta. The major difference between these descriptions of the nervous system is that Bouvier (1886) and I found no indications of zygoneury on either side of the nervous system, whereas Thiele (1928) found that the supraesophageal ganglion is con- nected to both the left and right pleural ganglia. Neither I nor Bouvier (1886) could locate the statocysts, but Thie- le (1928) described them next to the pedal ganglia. There appear to be several errors in Fretter & Graham’s (1994) comments regarding the nervous system of E. clathrus. They do not give a detailed description of the nervous system, but in their figure 99 they show the left cerebral ganglia located at the level of the jaws in a fully retracted proboscis, and they state that the nerve ring is at the level of the buccal ganglia in the retracted probos- cis. This is not the case in any of the epitoniids that have been described in more detail. In addition, the structure labeled as ‘‘Icg’’ in their figure is in the same location as the stylet bulbs in N. tincta, which superficially resemble ganglia. Conclusion Although adult anatomy clearly supports caenogastro- pod affinities, epitoniids possess developmental charac- ters that have previously been thought to characterize R. Collin, 2000 both heterobranchs and caenogastropods, monophyletic sister groups. Further careful homology assessment, more thorough taxon sampling, and detailed explicit cladistic analysis of gastropod relationships are necessary before the homologous and homoplasious characters can be identified with any certainty. This study highlights the fact that developmental characters that were previously thought to have good congruent distribution among gas- tropod taxa, consistent with higher level classification, may not in fact be so unambiguously distributed when more poorly known, basal, or intermediate taxa are ex- amined. Acknowledgments. 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Journal of the Marine Bi- ological Association of the United Kingdom 35:291-—305. The Veliger 43(4):313-318 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 The Ecology and Rapid Spread of the Terrestrial Slug Boettgerilla pallens in Europe with Reference to Its Recent Discovery in North America HEIKE REISE Staatliches Museum fiir Naturkunde Gorlitz, PF 300154, D-2806 Gorlitz, Germany (e-mail: Naturmuseum.GR.Reise. @t-online.de) JOHN M. C. HUTCHINSON School of Biological Sciences, University of Bristol, Woodland Road, Bristol, BS8 1UG, United Kingdom (e-mail: John.Hutchinson @bristol.ac.uk) ROBERT G. FORSYTH AnD TAMMERA J. FORSYTH 2574 Graham Street, Victoria, British Columbia, V8T 3Y7, Canada (e-mail: Robert-Forsyth @ bc.sympatico.ca) Abstract. The terrestrial slug Boettgerilla pallens Simroth, 1912, is reported from two sites on Vancouver Island, British Columbia, the first records for this Palaearctic species in America. This paper describes how to recognize the species, and summarizes European studies of its ecology. It is unusually wormlike in appearance, lives mostly under- ground, and occurs in a very wide range of habitats. This century the species has spread remarkably far and fast across Europe from the Caucasus. This is demonstrated by a table of first occurrences in each country, and by three case studies of spread within Great Britain, Belgium, and north-west Austria. We predict that it will spread rapidly in North America, and may already occur more widely, but there is no evidence that it will become an important pest. INTRODUCTION European slug species have been extremely successful colonists in other continents (Chichester & Getz, 1969; Rollo & Wellington, 1975; Barker, 1979, 1989). Arion subfuscus (Draparnaud, 1805), for instance, had already been discovered at several localities in eastern North America in the first half of the 19th century, and evidence points to repeated introduction events (Chichester & Getz, 1969). In many North American localities European slugs now predominate over native slugs, in terms of both abundance and number of species. Some are economi- cally important pests. However, some introduced species are usually still limited to synanthropic habitats. This paper reports the first finding in America of an- other Palaearctic slug species, Boettgerilla pallens Sim- roth, 1912. Within this century this species has spread from the Caucasus right across central and northwest Eu- rope, which suggests that the invasion in North America could rapidly become equally extensive. We hope here to draw workers’ attention to the possibility of this species occurring in their own neighborhood in the near future, if not already. Also workers in other continents should realize that it is a potential colonist. The literature on B. pallens that we have consulted is in several languages and spread over many journals. We consider here only the most pertinent aspects of its biol- ogy and cite only a selected minority of the publications dealing with the species, since most report little more than its discovery in a new location. We describe how to recognize B. pallens and where to look for it, summarize what is known of its ecology, and then discuss in greater detail its rate and pattern of spread. DISCOVERY IN BRITISH COLUMBIA We discovered B. pallens on 14 July 1998 at two sites at the southern end of Vancouver Island, British Columbia, Canada: from beside a trail below Denison Road, Wal- bran Park, east side of Gonzales Hill, Oak Bay District Municipality (48°024.7'N, 123°019.0’W); and in Centen- nial Park, near Graham Creek, Central Saanich District Municipality (48°035.2'N, 123°025.5'W). Specimens have been deposited in the Royal British Columbia Mu- seum, Victoria (catalogue number 998-00224-001) and in the Staatliches Museum fiir Naturkunde Gorlitz, Germany (catalogue numbers p5495 and p5496). The first locality is in a residential suburb adjacent to the city of Victoria, in scrubland at the back of gardens. The flora is a mixture of native and naturalized species, the latter presumably spread from the gardens. The sec- ond locality is 21 km away, in a rural area of suburban acreages and farmland. We found Boettgerilla pallens in a small wooded ravine dominated by Western Redcedar Page 314 (Thuja plicata), Douglas-fir (Pseudotsuga menziesii), and Bigleaf Maple (Acer macrophyllum). This site borders the developed portion of the park but is at least 100 m from the nearest habitation. At the first locality we found six individuals, and at the second four individuals, so the species appeared to be well established at these sites. All specimens were rough- ly half grown, which is compatible with the usual timing of the life cycle in Europe (see below). This discovery was preceded by a more widespread survey of non-marine Mollusca in British Columbia by RGF and TJE but this has concentrated on snails (For- syth, 1999). Subsequently, we specifically searched for slugs in several other synanthropic localities in Greater Victoria, in the city and suburbs of Vancouver, and else- where in mainland British Columbia, finding other intro- duced slugs but not B. pallens. However, as time was limited and the weather mostly rather dry, we cannot be confident that it does not occur more widely. DESCRIPTION Boettgerilla Simroth, 1910, is currently placed in its own family, Boettgerillidae Van Goethem, 1972. Three nom- inal species have been described—B. compressa Simroth, 1910, B. pallens, and B. vermiformis Wiktor, 1959—but the latter two names have been synonymized (Wiktor, 1972, 1973). Boettgerilla pallens is a small to medium-sized slug, up to 50 mm long when extended. When active it is un- usually slender and appears distinctively wormlike (Fig- ure 1). When contracted, it is of more normal proportions, and then the keel, which extends right up to the mantle edge, is more prominent. Full-grown animals are an un- speckled lead-grey color that fades to almost white in front of the mantle and on the lower flanks and sole. The sole may show a tinge of yellow, but the mucus is col- orless. Juveniles are paler, and very young slugs look al- bino. Young B. pallens, when contracted, might be con- fused with a young Deroceras, although their long keel is still distinctive. The pneumostome, which in the adult stands out as slightly paler than the mantle, is only a little more than halfway back along the mantle. Two grooves extend from the top of the pneumostome, one forward and one back, but these can be difficult to see after pres- ervation. Boettgerilla pallens has a thin internal shell, typically with an irregular outline. The genitalia are very distinct from those of other genera, particularly in respect of a spindle-shaped swelling along the vas deferens (e.g., see the figures in Van Goethem, 1972, or Colville et al., 1974; the latter includes a character for separating B. compressa). ECOLOGY Boettgerilla pallens occurs in a wide variety of habitats, including gardens, grassland, and both deciduous and co- The Veliger, Vol. 43, No. 4 niferous woodland. It is also tolerant of a wide range of soil types, and of soil water content, calcium content, and pH (3.2-7.8; De Wilde et al., 1983). It is predominantly subterranean, mostly found within about 25 cm of the surface, but sometimes at depths of up to 60 cm (Gunn, 1992; Seid] & Seidl, 1997). Wiktor (1973) further stated that it can burrow holes like an earthworm. Fortunately for malacologists, a small proportion of the population can also be found by searching under stones, under rot- ting wood, and in leaf litter. The species is strongly pho- tophobic, but a study using a rhizotron showed that the slugs underground are most active during the hours of daylight (Gunn, 1992). In North Wales mating and egg laying occurred below ground during late summer and into autumn (Gunn, 1992). The juveniles started to develop grey coloration from early May, and adults could survive until December. These observations are compatible with casual observa- tions from central Europe (Ant, 1966; Zeissler, 1981; Seid] & Seidl, 1997). However, Zeissler also proposed, on somewhat weak evidence, that in Romania possibly the development is delayed by hot dry summers, so that slugs would not become adult until the following spring. Gunn (1992) observed B. pallens feeding most often on earthworm feces, detritus, and soil surfaces. These slugs also ate decaying plant matter, fungal hyphae, car- rion, and living roots. This agrees with Daxl’s (1967) con- clusions that they prefer roots to green plant material (based on feeding preferences in the laboratory and gut dissections of wild-collected specimens). Other support- ing observations come from Schmid (1962), Zeissler (1964), Wiktor (1973), and von Proschwitz (1994). Boett- gerilla pallens has also been reported to eat slug eggs (Wiktor, 1961; Fechter & Falkner, 1990). There is no evidence that this species has ever become a horticultural or agricultural pest, but its subterranean habits might make this difficult to establish, especially as in Europe other slug species known to be pests are nor- mally also present. Individual B. pallens usually take only a few bites from any particular food item before moving on, which makes it less likely that the species would have a significant effect on root crops (Gunn, 1992; Seidl & Seidl, 1997). HISTORY oF SPREAD Boettgerilla pallens was first described from specimens collected in 1907 from natural habitats (shady montane forests) in the Caucasus, a mountain range forming the south-eastern border of Europe (Simroth, 1912). As this is also the only area where the sole other member of the family (B. compressa) is known, it is normally assumed that B. pallens has spread from there. Records of its oc- currence in natural habitats in this region all lie in Ab- khazia and western Georgia, on the south-western flank of the Great Caucasus (Likharev & Wiktor, 1980), but the H. Reise et al., 2000 Page 315 Figure | Adult specimens of Boettgerilla pallens from Europe. The specimen in the upper photograph (from Upper Lusatia, eastern Germany) is about 40 mm long. The lower photograph is a slightly dorsal view of a specimen freshly preserved in alcohol, where the scale has 1-mm divisions (collected from Leigh Delamere Motorway Services, Wiltshire, England). The keel here shows up most clearly where kinked near the tail, but it continues up to the mantle. areas around have probably been too little studied to be sure of the full extent of its natural range. In 1959 Wiktor described B. vermiformis (later syn- onymized with B. pallens) from Poland. This and his sub- sequent papers (Wiktor, 1960, 1961) drew attention to the possibility that it might be found elsewhere in Europe. There rapidly followed a rush of discoveries throughout much of Europe. Table 1 gives the dates of first collection in each country; 1949 is the earliest date outside the Cau- casus, based on preserved German material identified subsequently (Schmid, 1966). B. pallens has also now been reported from Turkey (Wiktor, 1994) and from other regions of the former USSR including Armenia, the Ukraine, the St Petersburg and Moscow regions of Rus- sia, and from well east of the Caucasus (in Tadzhikistan, and from Chelyabinsk in Western Siberia) (Likharev & Wiktor, 1980). The high rate of discovery in new countries in the 1960’s probably reflects the rate of spread of information rather than the speed of dispersal of the species. In order to judge how the species really spreads, we consider three cases where initially the species had not been recorded despite local malacologists being both actively involved in mapping and aware of this species’ existence. In Great Britain many malacologists contribute to the mapping scheme administered by the Conchological So- ciety of Great Britain and Ireland. The initial discovery of B. pallens in Britain in 1972 was well publicized in that society’s journals (e.g., Colville et al., 1974), and interest in mapping was also stimulated in the 1970’s by Page 316 Table 1 Date of first collection of Boettgerilla pallens in each country where it has been found (as defined before post- 1990 political changes, except for Ireland where the unit is the whole island). The references are not necessarily to the original publications, but do allow these to be traced. We have not found records for Italy, Greece, Bul- garia, Yugoslavia, Albania, Spain, or Portugal. Date of collec- Country tion Reference USSR 1907 Simroth, 1912 West Germany 1949 Schmid, 1966 Poland 1956 Wiktor, 1959 Czechoslovakia 1960 Schmid, 1963 East Germany 1960 Schmid, 1963 Switzerland 1960 Schmid, 1963 Belgium 1967 De Wilde et al., 1983 France 1968 von Proschwitz, 1994 Finland 1968 von Proschwitz, 1994 Romania 1969 von Proschwitz, 1994 Austria 1971 von Proschwitz, 1994 Hungary 1971 Varga, 1980 Great Britain 1972 Colville et al., 1974 Island of Ireland 1973 von Proschwitz, 1994 Netherlands 1973 von Proschwitz, 1994 Sweden 1974 von Proschwitz, 1994 Turkey 1985 Wiktor, 1994 Andorra 199] Borreda et al., 1996 Luxembourg 1996 K. Groh & G. Weitmann (pers. com.) Denmark 1998 von Proschwitz (pers. com.) Norway 1998 von Proschwitz (pers. com.) Canada 1998 this paper the publication of a distribution atlas (Kerney, 1976) and a popular guidebook (Kerney & Cameron, 1979). Figure 2a shows the British Isles divided into 153 vice-counties of roughly equal size and indicates the date of discovery of B. pallens in each. The species currently occurs in at least 91 vice-counties. Probably the data from Ireland and Scotland are too biased by intermittent recording effort to be interpreted, but the data in England and Wales should be more reliable. The pattern is of widely scattered discoveries, rather than a gradual spreading from a few nuclei. This suggests the importance of transport by man. After the initial discovery was publicized, it was not the case that many new sites were immediately reported; rather the number of vice-county records increased steadi- ly throughout the late 1970’s and the 1980’s (Figure 2b). The rate of increase has fallen sharply in the 1990’s. This cannot be explained by the reduced number of vice-coun- ties in which it remains to be discovered, but may be an artifact caused by recorders rarely visiting some vice- counties, by a general lowering of recording effort, or by The Veliger, Vol. 43, No. 4 40 w oS o 2 2 s i=} = ° 2 o A > 2 20 ° S o ao E 5 Zz 1975 1980 1985 1990 1995 Date Figure 2 The spread of Boettgerilla pallens in the British Isles as shown by new vice-county records. These have been published annually in the “Proceedings” section of the Journal of Conchology, and we add here Irish records given in Cawley (1998), and two re- cords from our own recent fieldwork. Kerney’s (1999) map of occurrences in 10-km grid squares implies that the species occurs in at least a further 10 British vice-counties, but these uncon- firmed records are not incorporated here. Dates given as, for instance, 1980-81 are counted as 1980. In Figure 2a the diameter of the dot correlates linearly with how long ago B. pallens has been known in that vice-county. For clarity, outlines of the vice- counties are shown straightened. The Channel Islands, and Ork- ney and Shetland are displaced in boxes. Figure 2b shows the increase in the cumulative number of these records for the 70 vice-counties in England and Wales. a reduced tendency to report what is now a less “‘excit- ing’”’ species (M. P. Kerney, personal communication). Our second case study is Belgium where B. pallens was first discovered in 1967. The evidence is good that it did not occur at least before 1950 (De Wilde et al., 1983). A project systematically mapping terrestrial mol- lusks on a 10-km grid started in 1970. From 1973 onward the data showed that B. pallens was widespread through most of Belgium except for the north. Although the north H. Reise et al., 2000 was extensively sampled in the 1970’s, only from 1981 onward did B. pallens appear there (De Wilde et al., 1983, 1986). In the south, although the localities already oc- curred over a wide area in the 1970’s, De Wilde et al. (1983) stressed that on a more local scale the species became more densely distributed, and spread to a wider range of habitats. The third case study is an extensive area in northwest Austria, which had been searched for slugs in 1994 and 1995 (Seidl & Seidl, 1997). Nevertheless, B. pallens was known from only 12 sites in this area until another search in 1997 which revealed 70 new localities in the same area. Seid] & Seidl concluded that the species was still spreading, especially because some of these new locali- ties had been searched several times earlier. All three case studies show rapid range expansion over a wide area, which suggests that dispersal was aided by human activities. This is supported by comments from several authors that the species was first found in syn- anthropic habitats or in more natural habitats adjacent to where garden waste had been dumped (e.g., Wiktor, 1973; De Wilde et al., 1983; von Proschwitz, 1994; Seid] & Seidl, 1997). It is easy to envisage that the subterranean habits of B. pallens facilitate its dispersal with garden plants distributed from commercial nurseries. The in- creasing distances that potted plants and root vegetables are transported probably explains why it is only in the last 50 years that it seems to have spread through Europe. However, B. pallens is also found in natural, or at least near-natural, habitats far away from settlements (e.g., Schmid, 1966; De Wilde et al., 1983; Seidl & Seidl, 1997). This has led some authors to question the as- sumption that the species is an introduction in central Europe (Ant, 1966; Schmid, 1966). But even in Britain, where it is clearly an introduction, it is now found com- monly in undisturbed habitats. So we presume that in cen- tral Europe the same process of colonization simply oc- curred at an earlier date before monitoring began (see also De Wilde et al., 1983; von Proschwitz, 1994). DISCUSSION What can this information from Europe suggest about the likely spread of B. pallens in North America? There are climatic and vegetational differences with Europe, but the species has shown itself very adaptable in these respects. Perhaps more important for a species that first colonizes synanthropic habitats are differences in human geogra- phy, such as in the distances between towns, the retailing of horticultural plants, or whether rubbish dumps occur adjacent to natural habitats. However, any such differenc- es have not stopped other European species from becom- ing common in North America, at least in urban areas (Chichester & Getz, 1969; Rollo & Wellington, 1975). Because the long-distance spread of B. pallens in Europe seems to have been much assisted by man, neither the Page 317 straits separating Vancouver Island from the mainland nor the Rocky Mountains look likely to be significant barriers to its dispersal eastward. What might prove more of a barrier is the border with the USA across which transport of soil and plants is restricted and actively controlled. If B. pallens succeeds in crossing this border, it is difficult to predict how much farther south it could spread before being limited by climatic factors. In Europe the species is not known from most of the countries to the south, bordering the Mediterranean, where the climate is gen- erally hotter (Table 1); this might partly be explained by less faunistic work in these countries, but at least in Spain, Italy, and Greece research on terrestrial mollusks is not undeveloped. It will be interesting to monitor how far and how quickly B. pallens spreads on Vancouver Island, to main- land British Columbia, and perhaps elsewhere. To do this it is most important to check now whether the species occurs already. In North America the occurrence of ter- restrial mollusks, and particularly of introduced terrestrial slugs, has been far less studied than in Europe. It is not at all improbable that B. pallens is already common in other parts of North America as a result of independent introductions from Europe. In Europe there is no evidence that B. pallens has been economically important. This will probably also hold true in North America, but there are fewer native slugs there, and maybe consequently fewer biological enemies of slugs, so its harmlessness cannot be guaranteed. Acknowledgments. We thank Professor Wiktor, Dr Kerney, and two referees for their helpful comments on the manuscript. Thanks also to Manfred and Louise for their generous hospitality while HR and JMCH collected around Vancouver. Wolfgang Jun- ius took the upper photograph in Figure 1, and Manfred Wanner kindly helped in its preparation. LITERATURE CITED ANT, H. 1966. Eine neue Nacktschnecke, Boettgerilla (pallens?) vermiformis, in Westfalen. Natur und Heimat 26:71-—74. BARKER, G. M. 1979. The introduced slugs of New Zealand (Gas- tropoda: Pulmonata). New Zealand Journal of Zoology 6: 411-437. BARKER, G. M. 1989. Slug problems in New Zealand pastoral agriculture. Pp. 59—68 in I. F Henderson (ed.), BCPC Mono- graph No. 41, Slugs and Snails in World Agriculture. British Crop Protection Council: Thornton Health, United Kingdom BorrepA, V., M. A CoLtapo, J. BLAsco & J. S. Espin. 1996. Slugs (Gastropoda, Pulmonata) of Andorra. Iberus 14:25— 38. CAwLey, M. 1998. A note on the status of the alien slug Boett- gerilla pallens Simroth (Mollusca: Gastropoda) in Ireland. The Irish Naturalists’ Journal 26:63. CHICHESTER, L. F & L. L. Getz. 1969. The zoogeography and ecology of arionid and limacid slugs introduced into north- eastern North America. Malacologia 7:313—346. COLVILLE, B., L. LLoyD-EvANs & A. Norris. 1974. Boettgerilla pallens Simroth, a new British species. Journal of Conchol- ogy 28:203-208. Dax, R. 1967. Ein Beitrag zur Biologie von Boettgerilla ver- Page 318 miformis Wiktor 1959. Zeitschrift fiir angewandte Zoologie 54:227-231. De WILDE, J. J., J. L. WAN GOETHEM, & R. MARQUET. 1983. Over de verspreiding, de uitbreiding en de oecologie van Boett- gerilla pallens Simroth, 1912 in Belgie. Studiedocumenten van het K.B.I.N., Brussel 12:1—31. DE WILDE, J. J., J. L. VAN GOETHEM & R. MARQUET. 1986. Dis- tribution and dispersal of Boettgerilla pallens Simroth 1912 in Belgium (Gastropoda, Pulmonata, Boettgerillidae). Pro- ceedings of the 8th International Malacological Congress, Budapest, 1983:63—68. FECHTER, R. & G. FALKNER. 1990. Weichtiere. Mosaik Verlag: Munich, Germany. 287 pp. ForsyTH, R. G. 1999. Distributions of nine new or little-known exotic land snails in British Columbia. The Canadian Field Naturalist 113:559-568. Gunn, A. 1992. The ecology of the introduced slug Boettgerilla pallens (Simroth) in North Wales. Journal of Molluscan Studies 58:449—453. KERNEY, M. P. (ED.). 1976. Atlas of the Non-Marine Mollusca of the British Isles. Institute of Terrestrial Ecology: Cambridge, United Kingdom v + 203 pp. KERNEY, M. P. 1999. Atlas of the Land and Freshwater Molluscs of Britain and Ireland. Harley Books. 261 pp. KERNEY, M. P. & R. A. D. CAMERON. 1979. A Field Guide to the Land Snails of Britain and North-West Europe. Collins: Lon- don, United Kingdom. 288 pp. LIKHAREV, I. M. & A. WIKTOR. 1980. The Fauna of Slugs of the USSR and Adjacent Countries (Gastropoda Terrestria Nuda). Fauna SSSR, Mollusca III (5), New Series 122. Akademii Nauk SSSR, Zoologicheskii Institut: Leningrad. 437 pp. [in Russian] PROSCHWITZ, T. VON. 1994. Boettgerilla pallens (Simroth)—a pe- culiar Caucasian slug species spreading in Sweden. Gote- borgs Naturhistoriska Museum, Arstryck 1994:45—54. Rotio, C. D. & W. G. WELLINGTON. 1975. Terrestrial slugs in the vicinity of Vancouver, British Columbia. The Nautilus 89:107-115. SCHMID, G. 1962. Die kaukasische Nacktschnecke Boettgerilla vermiformis auf dem Spitzberg bei Tiibingen. Natur und Mu- seum 92:263-—266. ScHMID, G. 1963. Zur Verbreitung und Anatomie der Gattung Boettgerilla. Archiv fiir Molluskenkunde 92:215—225. The Veliger, Vol. 43, No. 4 SCHMID, G. 1966. Weitere Funde von Boettgerilla vermiformis. Mitteilungen der Deutschen Malakozoologischen Gesell- schaft 1:131—136. SEIDL, FE & W. SEIDL. 1997. Die Wurmnacktschnecke Boettgerilla pallens Simroth 1912 im westlichen Ober@ésterreich und in den angrenzenden Gebieten. Mitteilungen der Zoologischen Gesellschaft zu Braunau 7:63—73. Smrorty, H. 1912. Neue Beitraége zur Kenntnis der kaukasischen Nacktschneckenfauna. Mitteilungen des Kaukasischen Mu- seums 6:1—140. VAN GOETHEM, J. 1972. Contribution a l’Etude de Boettgerilla vermiformis Wiktor, 1995 (Mollusca Pulmonata). Bulletin de l'Institut Royale des Sciences Naturelles de Belgique 48:1— 16. VarRGA, A. 1980. Ein neuer Fundort von Boettgerilla pallens Simroth 1912 in Ungarn. Soosiana 8:47—48. WIKTOR, A. 1959. Boettgerilla vermiformis n. sp. (Mollusca, Pul- monata). Communications of the Poznan Society of Friends of Science 4:1-2. WIkTOR, A. 1960. Description of Boettgerilla vermiformis Wiktor (Mollusca, Pulmonata). Bulletin de la Société des Amis des Sciences et des Lettres de Poznan (D) 1:151—156. WIkTorR, A. 1961. Materialien zur Kenntnis der Gattung Boett- gerilla Simroth, 1910 (Gastropoda, Limacidae). Annales Zoologici 19:125—145. WIKTOR, A. 1972. Boettgerilla pallens Simroth, 1919. Informa- tions de la Société Belge de Malacologie 1:145—148. WIkToR, A. 1973. Die Nacktschnecken Polens. Arionidae, Mil- acidae, Limacidae (Gastropoda, Stylommatophora). Mono- grafie Fauny Polski 1. Polish Academy of Sciences: Warsaw, Poland. 182 pp + figures. WIkTOoR, A. 1994. Contribution to the knowledge of the slugs of Turkey (Gastropoda terrestria nuda). Archiv fiir Mollusken- kunde 123:1—47. ZEISSLER, H. 1964. Boettgerilla vermiformis Wiktor, eine fiir Sachsen neue Nacktschnecke und ihre Begleitfauna (Mol- lusca, Pulmonata). Zoologische Abhandlungen und Berichte aus dem Staatlichen Museum fiir Tierkunde in Dresden 26: 277-280. ZEISSLER, H. 1981. Die alte Boettgerilla pallens-Fundstelle von Sinaia-Cumpatu (Rumanien) (Gastropoda, Stylommatopho- ra). Malakologische Abhandlungen des Staatlichen Muse- ums fiir Tierkunde Dresden 7:107—109. THE VELIGER © CMS, Inc., 2000 The Veliger 43(4):319-—329 (October 2, 2000) Successful and Unsuccessful Predation of the Gastropod Nucella lapillus (Muricidae) on the Mussel Mytilus edulis from Maine GREGORY P. DIETL Department of Zoology, North Carolina State University, Raleigh, North Carolina 27695-7617, USA Abstract. The behavioral components of prey selection of the muricid gastropod Nucella lapillus (Linnaeus, 1758) were reconstructed from 546 large (25-100 mm anteroposterior length), drilled valves of the blue mussel Mytilus edulis Linnaeus, 1758, collected from Sand Piper Beach, Appledore Island, Maine. Regression of outer borehole diameter on prey valve length indicates that muricids were selective of prey size, although the degree of correlation (7° = 0.117) is lower than for smaller size (< 35 mm) classes of mussels reported in the literature, indicating a relaxation of size selectivity by the predator in larger mussel size classes. The rare occurrence of boreholes in mussels up to 100 mm in length suggests that the relatively thin shell (~ 3 mm) of even large mussels alone was not an effective defense against drilling predators. Prey effectiveness in deterring drilling predators, indexed by the ratio of unsuccessful drilling attempts to the total number of attempted drillholes, increased as mussel valve length increased, suggesting a higher probability of interruption of the drilling process by abiotic or biotic factors later in ontogeny. Twenty-nine percent of boreholes initiated by the predator were either incomplete (n = 151) or repaired (n = 97) shortly after the drilling process was completed. In contrast to previously published results, drillholes are not preferentially clustered on the dorsal umbonal and posterodorsal portions of the valve. Cryptic placement of boreholes on the thicker ventral region of the valve may have reduced the risk of detection of the driller by other foraging predators during the prolonged drilling process of large mussels (> 4 days). INTRODUCTION The blue mussel Mytilus edulis Linnaeus, 1758 is a mod- erately large, epibyssate bivalve that is common along western Atlantic shorelines, attached to rocky substrates. A common predator of M. edulis is the intertidal to shal- low subtidal muricid gastropod Nucella lapillus (Linnae- us, 1758), which drills a cylindrical hole through the mus- sel’s shell (Figure 1) (see Carriker, 1969, 1981; Carriker & Van Zandt 1972; and Carriker & Williams 1978 for discussions of the drilling process). The stereotypic com- ponents of prey size- and drillhole site-selection by dog- whelks feeding on intertidal mussels have also been stud- ied extensively under both field and experimental condi- tions (Seed, 1969; Palmer, 1983, 1984; Hughes & Dun- kin, 1984a, b; Burrows & Hughes, 1989; Hughes & Drewett, 1985; Hughes et al., 1992; Hunt & Scheibling, 1998). Previous studies have concentrated on the predator- prey interaction among small upper shore populations of M. edulis (< 40 mm shell anteroposterior [A—P] length). But blue mussels may occasionally reach lengths in ex- cess of 100 mm (Stanley, 1970). It is reasonable to sus- pect that blue mussels may attain a valve length or thick- ness late in ontogeny that cannot be successfully handled by drilling muricids. Consequently, large and thickened valves of adult mussels may serve as effective morpho- logical defenses against drilling gastropods, as in other co-occurring predator-prey interactions (Vermeij, 1978; Palmer, 1983; Norberg & Tedengren, 1995; Elner, 1978; Hughes & Elner, 1979; Leonard et al., 1999). Drilling events are also not invariably successful (Fig- ure 1A—E K, L). Most studies have focused on successful drilling events and have not quantified the frequency of unsuccessful events. Incomplete boreholes are relatively common in muricid-mussel interactions (Vermeij, 1982; Seed, 1969) and represent either an interruption of the drilling process or a limit (thickness) to prey handling of the predator. The probability of interruption is propor- tional to drilling time (Hughes & Dunkin, 1984a); thus, large, thicker-shelled mussels may have accumulated multiple unsuccessful drillholes as the thickness (i.e., handling time) of the shell increased throughout ontoge- ny. Mussels have also been reported to repair boreholes that perforated the shell (Figure 1A, C—E L) (Seed, 1969; Griffiths & Blaine, 1994). Seed (1969:337), working with the Mytilus-Nucella interaction, stated that, ““where the whelk has been disturbed before doing any serious dam- age, the mussel can seal off the hole by means of a pearly concretion secreted by the mantle.”’ It is not known, how- ever, how frequently these repaired boreholes occur or in what size classes of M. edulis. This study reconstructs the foraging behavior of N. lapil- lus feeding on large mussels when the preferred smaller prey size classes are rare or absent. Alternative predatory Page 320 The Veliger, Vol. 43, No. 4 Figure | Predatory drillholes in valves of Mytilus edulis. A. Interior view of a valve with a blisterlike repaired borehole. The borehole penetrated the valve in the umbonal region of the shell. «1.0. B. Multiply drilled ventral portion of a valve; the posteriormost drillholes are repaired, and the anterior borehole is incomplete. <0.8. C. Anteriormost drillhole is incomplete, the next two are repaired, and the last three are complete. 0.8. D. Large blister in ventral portion of valve. Note complete borehole along the dorsal periphery of the repair. X1.0. E. Repaired drillhole in the dorsal portion of the valve and a more ventrally located complete borehole. 1.1. E Repaired drillhole. Note positions of anterior and posterior adductor muscle insertion (arrows) showing the absence of a nacreous shell layer. «1.0. G. An oversized complete borehole in the ventral-umbonal sector. 1.0. H. Multiple drillholes in the dorsal portion of the valve. Note location of the anteriormost drillhole along the dorsal valve edge. <0.6. I. Valve with four complete boreholes. Note the two juxtaposed ventral boreholes. 0.6. J. Three complete drillholes located in posteroventral region of the valve. Two of these drillholes are close to the valve edge. X0.7. K. Anteriormost drillhole is incomplete and the other is complete. X0.7. L. Interior view of a valve with two repaired drillholes. The anteriormost repair has flaked off, artificially producing a complete borehole. 1.0. strategies that exploit large prey with increased handling times are discussed and contrasted to predicted patterns of selective predation established under experimental condi- tions. MATERIALS ann METHODS Drilled valves of the mussel M. edulis were collected from a gravelly, shelly, high intertidal lag deposit on Sand Piper Beach, on the wave-protected side of Appledore Island, Maine. This sample affords the unique opportunity to investigate the predator-prey relationship between adult dogwhelks (> 25 mm in shell height; Hughes et al., 1992) and large M. edulis (> 35 mm A-P length); this relation- ship has not been quantified in the literature. Hunt & Scheibling (1998) reported that the outer diameter of the borehole (OBD) is positively related (7? = 0.986) to the height (H) of the shell of N. lapillus by the regression equation, OBD (mm) = 0.135 X H (mm)°°”’. This equa- G. P. Dietl, 2000 300 Frequency on (o) 0 20) 40 7 60° 80 100° 1120 A-P Length (mm) Figure 2 Size frequency distribution of anteroposterior (A—P) lengths of the valve of Mytilus edulis with complete boreholes. tion predicts that adult NV. /apillus greater than 30 mm in shell height drill boreholes greater than 1.0 mm in di- ameter. In this study, approximately 96% of complete drillholes have OBDs greater than 1.0 mm. The size-frequency distribution of valves of M. edulis with complete drillholes (Figure 2) suggests that this death assemblage was most likely a low-shore subtidal population of mussels. Even though smaller size classes (< 25 mm A-P length) may have been selectively win- nowed by currents, most low-shore subtidal populations of mussels are predominantly composed of only large mussels (Seed, 1969). Seed (1980) also stated that in low- shore habitats, sheltered from wave action, growth rates may be enhanced, resulting in increased size of mussels. Moreover, brachyuran crabs may influence the local dis- tribution and population structure of their mytilid prey, in low-shore and subtidal habitats, by preying selectively upon small mussels (Kitching et al., 1959; Ebling et al., 1964; Seed, 1990). Shell size was determined by measuring valve A—P length and dorsoventral height using Vernier calipers to the nearest 0.05 mm on 546 left and right valves of M. edulis. Both OBD and inner borehole diameter (IBD) were measured with a micrometer-calibrated binocular microscope. Outer borehole diameter and IBD are both reliable correlates of predator size (Carriker & Van Zandt, 1972; Hunt & Scheibling, 1998; Palmer, 1988). In order to evaluate selectivity of prey size, principal components analysis was used to describe covariation between the di- mensions of the borehole (predator size) and dimensions of the mussel valve (prey size) (Table 1). In addition, to facilitate comparisons with data on juvenile to subadult dogwhelk-mussel interactions (Hunt & Scheibling, 1998), OBD was regressed on mussel length to determine the Page 321 Table 1 Principal component analysis of the correlation matrix of dimensions of predator (OBD and IBD) and prey (Length and Height). Eigenvectors 1 D 3 4 Length 0.56 —0.43 0.00 0 7Al Height 0.57 —0.42 —0.11 0.70 OBD 0.46 0.52 0.72 0.05 IBD 0.39 0.61 —0.69 —0.06 Eigenvalues 2.24 1.43 0.27 0.06 Explained Variance 56.0 85", 6.8 eS flexibility (or amount of variation) in prey selection for different size classes of the predator (Figure 3). Seed (1969) and Hughes & Dunkin (1984a) have re- ported that N. lapillus commonly drills the dorsal regions in valves of M. edulis from exposed shores. Accordingly, the site of each complete drillhole was assigned to one of six sectors in a grid superimposed on the valve surface (Figure 4). The null hypothesis that all sectors were drilled equally was tested using a Chi-square goodness- of-fit test (Figure 5). Expected frequencies of boreholes in a sector were calculated to be proportional to the av- erage surface area of each sector. In this study, unsuccessful drilling attempts were sorted into two groups: (1) incomplete drillholes and (2) dril- lholes that perforated the shell but were later repaired by the mussel (Figure 1D, EK L). A Chi-square goodness-of- fit test determined if unsuccessful drillholes (¢ncomplete =I! 1b 7 to) 2.2415<0.001, N=555 10 20 30 40 50 60 70 80 90 100110 A-P Length (mm) Figure 3 Regression of outer borehole diameter of complete boreholes (OBD > 1.0 mm) on mussel anteroposterior (A—P) length for the sample of Mytilus edulis from Maine. Page 322 DORSAL fed c e) Oo = re Ww WW i - 2 : < O a VENTRAL Figure 4 Sector grid used to determine site location of the drillhole. and repaired types treated separately) were distributed among the six sectors of the valve in proportion to the observed frequency of complete drillholes for each sector (Table 2). Any differences in observed versus expected frequencies, of successful and unsuccessful drilling at- tempts, may reflect the extent of postero-anterior shifts of unsuccessful boreholes with post-drilling shell accretion. Higher observed relative to expected frequencies of un- successful drilling events in umbonal sectors (1—2) and lower than predicted observed frequencies in more central and posterior sectors (3—6) would suggest that postero- anterior shifts in borehole positions had occurred. A Poisson distribution was fit to the observed frequen- cies of boreholes for each group and a goodness-of-fit test (G-test) used to determine if the occurrence of a bore- hole on a valve was independent of the occurrence of other boreholes for multiply drilled valves (Table 3). A significant result would suggest that the probability of the occurrence of successive boreholes on a valve was not independent of the first drillhole. Poisson distributed data reflect the chance that the occurrence of multiple com- plete, incomplete, or repaired boreholes in any one valve is very small. Prey effectiveness (PE), the probability that an individ- ual survives in encounters with muricid predators, was calculated as the number of unsuccessful drilling attempts divided by the total number of attempted drillholes (Ver- meij et al., 1989) (Table 4). An effectiveness of 1.0 in- dicates that all drilling attempts by predators were unsuc- cessful, and conversely, an effectiveness of 0.0 indicates that the predator was always successful in encounters with its prey. RESULTS Anp DISCUSSION Association between Predator and Prey Size Principal components analysis: Muricid predator size is correlated with prey size. The loadings of the first eigen- vector (56.0% of variation) of the correlation matrix be- tween variables express covariation in the dimensions of The Veliger, Vol. 43, No. 4 Frequency Sector Figure 5 Plot of the observed (O) versus expected (@) number of com- plete boreholes per sector. Expected frequencies calculated pro- portional to average surface area. A chi-square goodness of fit test results in a nonrandom distribution (x? = 73.5; df = 5; P < 0.001). the drillhole (size of predator) and dimensions of the mus- sel valve (size of prey) (Table 1). The second eigenvector (35.7% of variation) expresses inverse variation in the dimensions of the mussel valve and drillhole. The size of the predator and the size of the prey are correlated. However, the distribution of drillholes sug- gests a predation system in which the prey do not obtain a size refuge from predation (i.e., adult N. lapillus (OBD > 1.0 mm) successfully prey on a wide size range of adult mussels). In this case, all adult mussels would be vulnerable to N. lapillus. Similarly, Dayton (1971) re- ported that even shells of the thicker, Mytilus californi- anus Conrad, 1837, greater than 80 mm, were commonly found with complete muricid drillholes. However, only twenty-two (4%) of the mussels with complete boreholes were larger than 70 mm in A-P length (Figure 2). Mus- sels up to approximately 55 mm are preyed upon by a wide range of predators (0.7—2.0 mm OBD), whereas larger mussel size classes (60—70 mm) are drilled by a narrower size range of predators (1.5—2.0 mm OBD). This pattern suggests that generally mussels greater than 70 mm in A-P length obtain a size refuge from all but the largest drilling predators. Linear regression: A comparison of the size selectivity between intertidal dogwhelk recruits (sensu early juve- niles, Hughes et al., 1992), post recruits (smaller juveniles to subadults), and adults in interactions with M. edulis indicates a relaxation of selectivity in the subtidal adult muricid-mussel interaction. Hunt & Scheibling (1998) have shown that prey size is highly correlated with pred- G. P. Dietl, 2000 Table 2 Contingency table for locations of complete (C), incom- plete (1), and repaired (R) boreholes according to six sec- tor grid (see Figure 4); comparisons treated separately. Chi-square goodness of fit results in a nonrandom distri- bution of boreholes (x? = 37.0 and 35.6; P < 0.001 in both cases). Observed Expected Sector frequencies Total frequencies complete and incomplete Cc I Cc I 79 4] 120 96 24 106 37 143 114 29 Total 593 149 742 593 149 complete and repaired Cc R Cc R 1 79 30 109 94 15 ator size (7° = 0.371, P = 0.001) for early juvenile dog- whelks and that the coefficient of determination (propor- tion of the variation in predator size determined by var- iation in prey size) decreases (7° = 0.254, P = 0.004) in larger dogwhelk size classes (small juveniles to sub- adults). The adult dogwhelk-mussel interaction of this study also shows a significant correlation between OBD (of adult dogwhelks) and mussel shell length (7? = 0.117, P < 0.001) (Figure 3). Adult muricids successfully pen- etrated a wider range of mussel size classes than did younger cohorts of dogwhelks. Numerous mismatches evident from the least-squares regression of OBD on prey length (Figure 3) indicate that many undersized muricids successfully penetrated unpredictably large mussels. Sim- ilarly, Griffiths & Blaine (1994) reported a breakdown in the relationship between predator and prey size for Nu- cella cingulata (Linnaeus, 1771), preying on large (mean 52 mm A-P length) Mytilus galloprovincialis Lamarck, 1819, from the west coast of South Africa. Constraint on size-selective foraging. A_ profitability curve, calculated as the ratio of predicted weight of tissue extracted to predicted handling times, for N. lapillus feed- ing on M. edulis is a monotonically increasing function of mussel shell length (up to 40 mm) (Hughes & Dunkin, 1984a). The profitability of mussels to N. lapillus declines as prey size increases beyond 40 mm, so that dogwhelks should selectively reject larger (> 40 mm) mussels en- countered during foraging bouts (Hughes & Dunkin, Page 323 Table 3 G-test for goodness of fit of observed frequencies of com- plete, incomplete, and repaired boreholes to expected fre- quencies based on a Poisson distribution. Expected fre- quencies for complete boreholes calculated using a truncated Poisson distribution. Expected frequencies less than five were pooled with an adjacent class to obtain a joint class with an expected frequency greater than five. Observed frequencies were also grouped to match the grouping of expected frequencies to facilitate calculation of the G-statistic. Number of Poisson boreholes Observed expected per valve frequencies frequencies Significance Complete boreholes (0) a wae! 1 417 409.6 G = 4.91 D} 66 77.1 Gadj = 4.90 3 11 9.7 df =3-2= 4 4 0.9 P < 0.05 Total 498 497.3 Incomplete boreholes 0) 425 413.4 1 97 115.3 G = 3.02 2 17 16.1 Gadj = 3.01 3 5) 1S df=3-2=1 4 2 0.1 P > 0.05 Total 546 546.4 Repaired boreholes 0 471 461.8 62 78.0 G = 3.10 2 10 6.6 Gadj = 3.09 3 2 0.4 df =3-2=1 4 1 0.02 P > 0.05 Total 546 546.8 1984a). A correlation between predator and prey size is consistent with an energy-maximizing foraging strategy (Hughes & Burrows, 1990; Hughes et al., 1992). Dog- whelks are expected to select larger prey, the preferred size increasing with increasing predator size. In the present study, however, the dogwhelk population may not have had the more profitable smaller (sensu in- termediate of Hughes & Dunkin, 1984a) mussel size clas- ses to prey selectively upon. Furthermore, adult dog- whelks are capable of successfully drilling a wide range of potential prey given sufficient time to complete the drilling process (evident from the low correlation between predator and prey size, 7? = 0.117) (Figure 3). Thus the risk of attacking a larger mussel, which may be devalued by the predator because of the associated longer handling time, may have been unavoidable if more profitable prey size classes were unavailable or infrequently encountered. This foraging strategy was also documented for N. cin- gulata feeding on M. galloprovincialis by Griffiths & Blaine (1994). The most profitable prey when encoun- Page 324 Table 4 Prey effectiveness (PE) in deterring predation by drilling for M. edulis (PE = number of incomplete and repaired drillholes/total attempted drillholes). Overall PE = 0.29. Size class (mm) Complete Incomplete Repaired PE 20-30 11 1 0) 0.08 30—40 122 20 5 0.17 40-50 254 46 25 0.22 50—60 125 40 15 0.36 60-70 62 33 36 0.53 70-80 22 9 12 0.49 80—90 1 2 3 0.83 90-100 (0) 0) 1 1.0 tered are selected, but if relative abundance or encounter rates with the most profitable prey are low, less profitable prey are selected by the predator (Hughes, 1980). Thus selective feeding of N. lapillus is constrained in mussel beds consisting of predominately large prey (> 40 mm). Site Stereotypy in Positioning of the Borehole Complete boreholes: Borehole site stereotypy is evident for N. lapillus, although all sectors host complete bore- holes. Sectors 1 and 2, the umbonal regions of the shell, were preferentially drilled (x? = 73.5; P < 0.001) (Fig- ures 1G, 5). Drillholes in sector six, the posterodorsal region of the valve (Figure 1J), were less frequent than expected based on the surface area of the sector (Figure 5). Positioning of the borehole by the predator in the an- terior region of the valve (sector 2) is consistent with the results obtained by Seed (1969) and Hughes & Dunkin (1984a). According to Seed (1969:340), placement of the borehole in the umbonal and posterodorsal regions is pre- ferred, because “‘the shell is particularly thin and brittle either through excessive erosion (especially at the anterior end) or the absence of nacreous shell in the regions of muscle insertion (especially around the region of the pos- terior adductor muscle).”” (See Figure 1F for position of the anterior and posterior adductor muscles.) Surprisingly, approximately fifty percent of complete boreholes in subtidal mussels were positioned in the ven- tral sectors of the valve (Table 3) (Figure 1B, C, G, J, K). Selection of ventral drilling sites is confounding be- cause of the increased handling times associated with drilling through the thicker valve wall in this region of the shell (Hughes & Dunkin, 1984a). In contrast, Seed (1969:340) noted that complete drillholes were dispro- portionately concentrated on the dorsal portion of the valve of a sample of intertidal M. edulis (only an esti- mated 15% of drillholes occurred in ventral sectors). This contrast in the placement of the drillhole in mussel prey The Veliger, Vol. 43, No. 4 by N. lapillus may reflect a trade-off between foraging and the risk of predation, due to differences in the abun- dance and diversity of exposed versus submerged dog- whelk predators. To an individual dogwhelk the risk of predation may depend not only on the abundance of its predators, but also its own behavior. Foraging by the dogwhelk may increase the probability of mortality to predation (Gil- liam, 1990). A shift in prey size-selectivity from large to small prey may reduce the time the forager is vulnerable, to predation, thus decreasing the probability of death to predation. Indeed, Palmer (1984) suggested that drilling muricid gastropods from the west coast of North America select small prey to reduce the risk of predation and other attendant risks of handling prey with long subjugation times. But, the predator-mediated shifts in prey size-se- lectivity may be constrained by the absence of smaller mussels. Concentration of boreholes in the ventral sectors sug- gests a more inconspicuous drilling posture of the dog- whelk that is potentially less vulnerable to locally com- mon crabs such as Carcinus maenas Linnaeus, 1758, and Cancer irroratus Say, 1817, (Elner, 1978; Hughes & El- ner, 1979; Moody & Steneck, 1993), as opposed to being perched up and consequently more exposed on the dorsal margin of the shell. The risk of being attacked by mol- luscivores for N. lapillus may favor a suboptimal drilling site in the absence of alternative prey. The ventral dril- lhole location suggests a more cryptic drilling position that may provide a refuge from foraging predators during the increased handling times imposed by larger prey for N. lapillus in the absence of small mussels. If dogwhelks preferentially drilled the thicker ventral sectors rather than the more easily penetrated dorsal re- gions of the shell to reduce the risk of being detected by predators, then the proportion of unsuccessful events should be higher in the dorsal sectors (due to an increased likelihood of disturbance). However, 67% of incomplete boreholes are located in the ventral sectors of the valve (x? = 14.33; P < 0.001). Moreover, 36 and 21% of all boreholes in the ventral and dorsal, respectively, portions of the valve are unsuccessful drilling events (either in- complete or repaired). This pattern seems inconsistent with the selection of a cryptic drilling site to reduce the risk of predation. But a more ventral drilling position, especially within sector 3 (Figure 4), also increases the likelihood that the drilling process will be interrupted by the adhesion of byssi by the mussel to the shell of the drilling predator (Davenport et al., 1996; Wayne, 1987). Interruption of the long drilling process of a snail posi- tioned on the more exposed dorsal margin of the shell may result in death for the drilling predator (dogwhelks up to ~ 35 mm in shell height are vulnerable to crab predation; Hughes & Elner, 1979). In contrast, interrup- tion of a snail drilling in the more cryptic ventral sectors G. P. Dietl, 2000 may only result in temporary entrapment in a mussel’s byssus. Incomplete boreholes: Both incomplete and repaired boreholes accumulated in the ventral umbonal sectors in greater frequencies than expected (x? = 37.0 and 35.6, P < 0.001, respectively) (Table 2). The number of incom- plete boreholes in umbonal sector | is nearly twice the expected number (41 versus 24) (Table 2). Repaired bore- holes show a similar distribution for sector 1 (30 versus 15). The low numbers of both incomplete and repaired drillholes in the more posterior sector 4 relative to ex- pected frequencies (Table 2) suggest that postero-anterior shifts influenced the observed distribution of unsuccessful events recorded on the shell at death. Unsuccessful at- tempts to drill the posterior margin of a young adult would leave an incomplete borehole that would occupy an increasingly more central and ultimately an anterior position late in ontogeny. Relationship between Thickness of the Valve and Incomplete Boreholes Carriker & Van Zandt (1972) have suggested that the muricid Urosalpinx cinerea (Say, 1822), selects a drilling site without regard to thickness of the shell (but see dis- cussion by Hughes & Dunkin, 1984a for N. lapillus prey- ing on mussels). Maximum depth of penetration is limited by the length of the fully extended proboscis of the pred- ator. Proboscis length in N. lapillus approximates dog- whelk shell height (Hughes & Dunkin, 1984a). Given that shell thickness is phenotypically labile in M. edulis, being generally thicker in geographic areas of high predation pressure (Leonard et al., 1999), incomplete boreholes may represent a limit (i.e., thickness) to predation for WN. lapillus feeding on large mussels. Thickness of the mussel valve in the Nucella-Mytilus interaction did not limit predation for the size range of predators sampled (0.7—2.0 mm OBD). Valve thickness remains relatively thin, approximately 2.2 mm for even an 80 mm mussel in the umbonal region of the shell (es- timated from regression equations given in Hughes & Dunkin, (1984a)). Moreover, based on shell thickness, any given size class of predator (> 1.0 mm OBD) was capable of drilling a mussel 70 mm in A-P length (Figure 3). Given an average drilling rate of 0.36 mm day’, for N. lapillus drilling M. edulis, a shell 2.0 mm thick would take about 5.5 days to penetrate (Hughes & Dunkin, 1984a). Thus incomplete boreholes most likely represent interruptions of the drilling process. Frequency of Multiple Boreholes Complete boreholes: The goodness-of-fit (G-test) of the observed frequencies to those expected by the Poisson distribution indicates that multiple complete boreholes (Figure 1H—J) occur in greater frequencies than predicted Page 325 (Gadj = 4.91; P < 0.05) (Table 3). The clumped fre- quencies of complete boreholes in the tail of the distri- bution indicate that the occurrences of multiple boreholes on a single valve are not independent of each other. The occurrence of one successful drilling event enhances the probability of a second event occurring. Poisson proba- bilities of the occurrence of multiple complete boreholes on a single valve decrease as the number of complete boreholes on a valve increases (Figure 6), suggesting that multiply drilled valves are considerably rarer than singly drilled valves. This clumping of multiple complete boreholes may oc- cur due to the near synchronous initiation of the drilling process by conspecifics of N. lapillus on the same prey. Hughes & Dunkin (1984a) reported that only 60% of mussel tissue is extracted from 40 mm-long prey, 1.e., the point of satiation for a given snail will occur before the entire mussel has been consumed. Thus, aggregations of conspecifics on the same mussel shell may continue the drilling process, penetrating the shell at different times (or simultaneously?). The maximum number of snails that can mount and successfully drill the same mussel concurrently (produc- ing multiple complete drillholes) may depend upon the interval of time between initiation of drilling by different snails. Brown & Alexander (1994) reported that the te- nacity of Stramonita haemastoma (Gray, 1894), feeding on Crassostrea virginica (Gmelin, 1791), from the Gulf of Mexico was not dependent on the number of conspe- cifics that ‘‘joined”’ in the attack, but rather on the timing of the attacks. Multiple drillholes were observed in oyster valves, which suggested that most snails were attracted to the prey early and attempted to drill a hole. However, late-arriving snails may have abandoned unfinished dril- lholes to feed through the gaped valves. These snails were “cheating,” sharing in the rewards but not the cost of foraging. The valves of M. edulis also gape after pro- longed feeding of a drilling predator (Hughes & Dunkin, 1984a); thus it is reasonable to assume that some dog- whelks would abandon unfinished drillholes, although field observations are needed to substantiate this hypoth- esis. Similarly, Carriker & Van Zandt (1972:234) reported this behavior in U. cinerea feeding on C. virginica stat- ing, “‘as feeding by the snail(s) continues, the adductor muscle of the oyster weakens and relaxes, and the valves begin to gape slightly’’; subsequently, other conspecifics join in the attack by feeding through the gape between the valves. However, it should be noted that the frequency of valves with multiple complete boreholes (Table 4) sug- gests that this behavior is either unlikely for or ineffec- tually practiced by WN. lapillus feeding on large M. edulis, because as the first snail finished the drilling process, the other snails continued the slow process of excavating a drillhole, rather than adopting the more efficient ‘‘cheat- ing”’ strategy. Studies are needed to determine how long oS in 0.25 Poisson Probability Number of Boreholes Figure 6 Plot of calculated Poisson probabilities of the occurrence of mul- tiply complete (O), incomplete (LJ), and repaired (© ) boreholes in a single valve. the valves of M. edulis remain adducted after the initial dogwhelk(s) penetrate the shell. Repaired boreholes: The distribution of multiple re- paired boreholes (Figure 1L), unlike the distribution of multiple complete boreholes, followed a Poisson distri- bution, indicating that the probability of surviving a suc- cessful penetration of the valve was independent (Gadj = 3.09; P > 0.05) of the number of encounters with drilling dogwhelks. Therefore, the majority of multiple repaired drillholes on a single valve can be inferred to have oc- curred independently of each other (i.e., resulting from successive attacks by individual predators rather than multiple snails drilling concurrently). Indeed, Griffiths & Blaine (1994) suggested that large M. galloprovincialis accumulate multiple sublethal attacks (predatory ‘‘crop- ping’) by N. cingulata over time. Concurrent attacks by more than one snail would most likely result in the mus- sel’s death. This ability to repair perforations in the shell away from the mantle edge shortly after dislodgement of the mounted predator is advantageous because any breach in the valve may allow chemical cues to attract other drilling snails, durophagous crabs, or asteriids (Vermeij, 1983; Carriker & Van Zandt, 1972; Elner, 1978; Hughes & Elner, 1979; Moody & Steneck, 1993; Reimer & Te- dedgren, 1996), which are common subtidal coinhabi- tants. Indeed, the calculated Poisson probabilities (Figure 6) for the occurrence of multiple repaired drillholes are lower in comparison with probabilities for multiple com- plete and incomplete boreholes occurring on the same valve. The repair of perforations in the shell of M. edulis re- quires up to 6 to 8 weeks in water temperatures of 18°C The Veliger, Vol. 43, No. 4 (Meenakshi et al., 1973). (This slow experimental rate of repair may not reflect field conditions. Indeed, for 25—40 mm M. edulis, perforations < 2.0 mm in diameter are often repaired within a week in water temperatures of 21°C [R. R. Alexander, personal communication, 1999].) Consequently, there is considerable time for the injured mussel to be detected by predators such as C. maenas (Vermeij, 1983). The concentration of repaired drillholes in the larger size classes (mean mussel length = 59.4 and 47.9 mm for valves with repaired and complete bore- holes, respectively, t = 9.66, P < 0.0001) may have been generated by the fact that at sizes greater than 50 mm in A-—P length, mussels are effectively immune from attack by manipulative chelate predators such as C. maenas (EI- ner & Hughes, 1978; Seed & Hughes, 1995; Ameyaw- Akumfi & Hughes, 1987). Smaller mussels may physio- logically be able to repair perforations in the shell, but because they are in the size range vulnerable to foraging crabs, they are more prone to attack during the prolonged repair phase, and therefore less likely to show repaired injuries. Indeed, Seed (1969), working with intertidal mussels < 50 mm in A-P length, rarely encountered re- paired drillholes. In this study, only 5.2% of the total number of repaired boreholes were apportioned in the 20—40 mm size classes (Table 4). Similarly, no repaired drillholes were found in shells of M. galloprovincialis less than 40 mm (Griffiths & Blaine, 1994). Alternatively, Griffiths & Blaine (1994:fig. 3; p. 348) have suggested that there is a critical ratio of prey size to predator size above which prey survivorship is high in predator-prey interactions involving N. cingulata and M. galloprovin- cialis. Below this threshold, prey were almost always killed and completely consumed. Prey Effectiveness in Deterring Drilling Predators Effectiveness and adaptation: The effectiveness of an individual in a given environment is the probability that the individual survives in an encounter with a potentially lethal hazard, e.g., a predator (Vermeij, 1982a). An ef- fectiveness of 0.0 suggests that the predator was always successful. To recognize a character or behavior that con- fers an adaptive advantage, individuals must survive in encounters with predators (Vermeij, 1982a). Prey effectiveness (PE) in deterring drilling muricid gastropods was 0.29. Vermeij et al. (1989) reported a val- ue of 0.17 for a geographically proximal sample from Sheep Island, off Deer Isle, Maine. Prey effectiveness es- timates increase as shell size increases (Table 4). This pattern suggests that dogwhelks were less efficient in dealing with increasingly larger and thicker shells. How- ever, the accumulation of unsuccessful drilling attempts with size most likely reflects an increased probability of encountering drilling muricids over the lifespan of an in- dividual. Indeed, the distribution of multiple incomplete boreholes followed a Poisson distribution indicating that G. P. Dietl, 2000 the majority of incomplete boreholes occurred indepen- dently of each other as the result of successive attacks by individual predators (Table 3). Clearly, as the lifespan of the mussel increases, the likelihood of encountering a drilling predator also increases. From the point of view of the surviving prey there may not be a difference between an incomplete and a repaired drillhole (G. J. Vermeij, personal communication 1998), i.e., the mussel survived both the drilling event that cul- minated in an incomplete borehole and that which pro- duced a complete hole that was later repaired. The ability to repair complete boreholes, however, may be a physi- ological adaptation that increases the survivorship of the prey. Repaired boreholes represent survival of the sub- jugation phase of a predation event, a necessary condition for the recognition of adaptations that confer benefits with respect to survival (Vermeij, 1982). Whether or not the cessation of the drilling (subjuga- tion) phase of predation was due to interruption of the esurient predator by extrinsic biotic or abiotic factors, or to an intrinsic active defensive behavioral response of the mussel, is not known. Dogwhelks are actively entrapped by adhesion of byssi to the snail’s shell, which usually only temporarily immobilizes the snail (Davenport et al., 1996; Wayne, 1987; Carriker, 1981). In addition, Wayne (1987) has described behavioral interactions between M. edulis and Nucella emarginata (Deshayes, 1839), and Nu- cella lamellosa (Gmelin, 1791). After initial contact with the dogwhelk, the mussel’s valves alternately gaped wide- ly (retracting the mantle) and closed (this behavior may pinch the muricid’s foot causing it to move away from the valve edges). Indeed, only a few valves had complete boreholes near the valve margins (Figure 1J). Further- more, large mussels may be able to dislodge mounted snails by a sweeping motion of the narrow, fingerlike foot. Wayne (1987) described a similar action of the foot as “‘probing and wiping.’’ Contact with the mussel’s foot elicited a “‘lifting and twisting’? behavior in the dog- whelk, as well as directed retaliatory rasps at the foot of the mussel with the radula (see Wayne, 1987:fig. 1, p. 141). The ability to repair drillholes is an adaptation that complements the high probability of disturbance of dril- ling predators in this shallow-subtidal habitat. Sealing off a perforation in the valve would decrease the probability of detection by other predators, thus increasing survivor- ship. The variegated behavioral defenses (as well as in- ducible morphological defenses; i.e., thickness (Leonard et al., 1999)) of M. edulis function to increase the like- lihood of successfully deterring attacks by specialized predators (i.e., drilling muricids), thus increasing fitness by reducing the risk of predation. The valve flapping movements, flailing foot extensions, and byssus attach- ment by M. edulis described by Wayne (1987) not only act to deter drilling predators upon initial contact, but may also increase the chances of disturbing muricids that Page 327 have penetrated the interior of the shell with the exten- sible proboscis. Studies documenting the effectiveness of mytilids in different phases of predation by drilling mur- icid predators are needed to independently test whether the inferred defensive adaptations of extant species have evolved specifically in response to drilling muricids. Are active behavioral defenses more successful after the pred- ator completes the drillhole? Bias in prey effectiveness?: If N. lapillus actively aban- dons an unfinished drillhole in favor of feeding through the gaping valves in prey that had already been success- fully penetrated by conspecifics, then PE estimates may overestimate the defensive value of the mussel’s shell. One hundred valves in the sample contain both complete and incomplete boreholes. Incomplete boreholes found in association with complete boreholes may either represent a “‘true”’ unsuccessful drilling attempt most likely due to interruption of the prolonged drilling process of large M. edulis, or, alternatively, an incomplete borehole may rep- resent a muricid that ‘“‘cheated’”’ and abandoned a drillhole in foraging bouts where snails initiate the drilling process concurrently. The latter situation will artificially inflate the PE estimate. The predatory attack of several snails (sensu group foraging of Brown & Alexander, 1994) may result in multiple complete and incomplete boreholes in the prey’s valve. In these cases, however, the mussel did not survive the attack even though the shell recorded the abandoned incomplete drillhole as an unsuccessful dril- ling event. In contrast, repaired drillholes and valves with only incomplete drillhole(s) represent unquestionable sur- vival of an unsuccessful drilling event. Only 47 (8.6%) valves had evidence of unsuccessful (incomplete and re- paired) drilling events recorded on the valve not in as- sociation with the successful events. Another important caveat to the estimation of PE in deterring muricid drilling predators is the possibility of the recognition and reoccupation of failed drilling at- tempts by other muricid conspecifics. Resumed drilling of a formerly incomplete borehole not only decreases the handling time of the prey, but also the calculated PE es- timate. Reoccupying incomplete boreholes will decrease the number of unsuccessful drilling events recorded on the shell (decreasing the numerator in the PE ratio); thus PE estimates may also be slightly underestimated. This behavior of reselecting formerly initiated drillholes has been documented for N. lapillus feeding on M. edulis (Hughes & Dunkin, 1984a). Concluding Remark This study illustrates how much is still not known about the predator-prey relationship between N. lapillus and large size classes of M. edulis. Clearly, there is a need for experimental work to determine the cost and benefits of the reconstructed foraging behaviors of N. lapillus feeding on large mussels when the preferred smaller prey Page 328 size classes are rare or absent. Constraints on foraging may include the increased probability of mortality to pre- dation with increased handling time of large prey and active behavioral defenses of the prey which disrupt the drilling process. Alternative predatory strategies (i.e., se- lecting a ventral drilling position, group foraging, and predatory ‘‘cropping’’(?)) that exploit large prey with in- creased handling times are also observed in other pred- ator-mussel interactions (see Ameyaw-Akumfi & Hughes [1987]). Similar foraging ‘‘rules’”” may also apply for the adult N. lapillus-M. edulis predator-prey interaction. 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Traces and trends of predation, with special reference to bivalved animals. Palaeontology 26:455—465. VERMEU, G. J., E. C. DUDLEY & E. ZIpSER. 1989. Successful and unsuccessful drilling predation in Recent pelecypods. The Veliger 32:266—273. WAYNE, T. A. 1987. Responses of a mussel to shell-boring snails: defensive behavior in Mytilus edulis?. The Veliger 30:138-— 147. The Veliger 43(4):330—337 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 Predation on Juvenile Aplysia parvula and Other Small Anaspidean, Ascoglossan, and Nudibranch Gastropods by Pycnogonids C. N. ROGERS,* R. DE NYS AnpD P. D. STEINBERG School of Biological Science, University of New South Wales, Sydney 2052, NSW, Australia Abstract. This study investigates the predation on opisthobranchs by the pycnogonid Anoplodactylus evansi. Ano- plodactylus evansi attacked and fed on small individuals of 13 different species of anaspidean, ascoglossan (sacoglossan), and nudibranch gastropods in this study. This predator was used to investigate the feeding deterrent effects of diet- derived compounds in the sea hare Aplysia parvula. Anoplodactylus evansi was not deterred from feeding on A. parvula containing diet-derived secondary metabolites from the red seaweeds Laurencia obtusa or Delisea pulchra. However, increasing the quantity of metabolites present in A. parvula by treatment with adult extracts did deter A. evansi from feeding, suggesting that high levels of diet-derived materials in sea hares may have deterrent effects against a predator. A. evansi does not appear to sequester algal secondary metabolites from its prey. The abundance of A. evansi and A. parvula on the alga L. obtusa was measured to determine if this pycnogonid influences the population of sea hares in the field. A. parvula occurred in significantly lower abundance on L. obtusa thalli inhabited by A. evansi over consecutive years, a pattern consistent with population-level effects by this predator. INTRODUCTION Opisthobranch mollusks feed on a wide range of both macrophytes and animals, although most species have di- ets limited to a few prey types (Thompson et al., 1982; Paul & Pennings, 1991; Trowbridge, 1991; Rogers et al., 1995). Opisthobranchs may deter potential predators by storing noxious materials in their tissues. Such materials include chemical compounds or physical structures, which are obtained from dietary sources or produced by the animals de novo (Faulkner & Ghiselin, 1983; Karuso, 1987; Avila, 1995). A predator attempting to consume different species of opisthobranchs is likely to encounter a variety of defensive materials which it must overcome in order to successfully feed. The trophic flexibility re- quired to feed on different opisthobranchs is thus proba- bly high, and there are few reported generalist predators of these mollusks. Exceptions are some aglajid cephal- aspideans which prey on other opisthobranchs (Kandel, 1979; Pennings, 1990a; Faulkner, 1992). Other predators of opisthobranchs include anemones (Nolen et al., 1995), crabs (Carefoot, 1987; Trowbridge, 1994), fishes (Pen- nings, 1990a; Trowbridge, 1994), seastars (Carefoot, 1987), a pycnogonid (Piel, 1991), and birds (Todd, 1981). Ecological studies of predation on opisthobranchs have found that predator success varies, depending on opistho- branch body size (Pennings, 1990b), habitat complexity (Pennings, 1990b), and the presence of chemical defenses (Hay et al., 1989). Studies on the effects of acquired di- * Corresponding author: Phone: 61 02 9385 3738, Fax: 61 02 9385 1558, e-mail: Cary.Rogers@unsw.edu.au etary compounds as defenses have mainly investigated palatability to “‘potential’’ predators. Few studies have investigated the effects on predators of different types or quantities of compounds stored by opisthobranchs. One exception is the study on sea hares by Pennings (1990b) who found that chemically depauperate Aplysia califor- nica Cooper, 1863, was rejected less frequently by fishes than chemically rich conspecifics. The types and levels of defensive compounds stored by opisthobranchs may be important in determining a predator’s response follow- ing contact, and so influence the outcome of an attack. Population-level effects of predation on opisthobranchs have rarely been addressed. Sarver (1979) found high rates of mortality for juvenile Aplysia juliana Quoy & Gaimard, 1832, and attributed this to both abiotic and biotic factors including predators. Trowbridge (1994) used cages to exclude predators from mat-forming algae inhabited by ascoglossans in the field and found a signif- icant difference in the abundance of the ascoglossan AI- deria modesta Lovén, 1844, between caged and uncaged areas after 12 days. These studies suggest that rates of predation on some opisthobranchs may be high in the field, especially for small species or juveniles. In contrast, many adult opisthobranchs are generally regarded as free from predation with few reported natural enemies (Care- foot, 1987; Faulkner, 1992). This study investigates predation on opisthobranchs by the pycnogonid Anoplodactylus evansi Clark, 1963. Is- sues addressed are: (1) Where does A. evansi occur relative to opistho- branchs on a local scale? C. N. Rogers et al., 2000 (2) What is the diet breadth of A. evansi across different sizes and species of opisthobranchs? (3) How does the type and quantity of algal secondary metabolites stored by Aplysia parvula Guilding in Morch, 1863, affect predation by this pycnogonid species? (4) What is the abundance of A. evansi relative to the abundance of the sea hare A. parvula on the red sea- weed Laurencia obtusa Lamouroux, 1813? MATERIALS anpD METHODS Study Sites and Collection of Study Organisms Collection and observation of the pycnogonid Anoplo- dactylus evansi, the sea hare Aplysia parvula, and the red alga Laurencia obtusa were done in Port Jackson, near Sydney, New South Wales (NSW), Australia, which is a warm temperate region. Four sites where L. obtusa oc- curred were used in this study (Figure 1). All four sites had sandstone reefs covered by mixed algal assemblages dominated by brown macrophytes (described by Farrant & King, 1982). Aplysia parvula was also collected from the red alga Delisea pulchra Greville, 1830, at Bare Is- land. The other species of opisthobranch used in this study (Table 1) were collected from the algal beds or sponge gardens on the reefs surrounding Bare Island (de- scribed by Van der Velde & King, 1984), near the en- trance to Botany Bay, just south of Port Jackson. Subtidal collection of pycnogonids and opisthobranchs for exper- iments was done by searching algae, sponges, or other encrusting organisms in situ, removing the animals by hand, and placing them in a plastic container for trans- portation to the laboratory. Animals were held in the lab- oratory for observation and experiments in 10 L plastic aquaria supplied with flowing seawater, with pieces of algae (20—40 g wet weight [ww]) provided as habitat. Anoplodactylus evansi is reported to occur at several lo- calities in south eastern Australia (Clark, 1963). However, this pycnogonid broods its young and does not swim, and consequently its dispersal ability is limited to movement between adjacent algae and perhaps rafting (e.g., on dis- lodged algae) to new habitats. All pycnogonids used in experiments were non-egg-bearing adults (leg span 20-— 30 mm). A voucher specimen of the pycnogonid A. evan- si was placed with the Australian Museum in Sydney, accession number AM-P49749, the specimen being an egg-bearing adult male. Pycnogonid Feeding To determine the diet breadth of Anoplodactylus evan- si, 13 species of anaspidean, ascoglossan, and nudibranch opisthobranchs, and other small invertebrates were placed in contact with adult pycnogonids held in aquaria. Opis- thobranchs in a range of sizes from 50 to 820 mg (ww) were used (approximately 5—40 mm body length); the Page 331 other invertebrates tested were 5-15 mm in length. Test animals were observed to determine if they were attacked by pycnogonids, and if feeding commenced. Aquaria were checked on subsequent days to determine if prey items were consumed. The method of attack and con- sumption by A. evansi was observed in aquaria, and using a dissecting microscope for prey items isolated in a glass dish. When sea hares were attacked, it was noted if ink or opaline secretion was released, and the effect this se- cretion had on pycnogonid behavior. Once pycnogonids had ceased feeding on sea hares, any remaining body parts were identified. To investigate more rigorously how the capture success of pycnogonids related to prey size, individual pycnogo- nids and Aplysia parvula were weighed (ww) and placed in experimental compartments with a piece of insect mesh (1 mm) as a substratum. Predation trials were done in 25 L experimental aquaria which were partitioned into 20 separate compartments by plastic sheets. Seawater flowed through the compartments via insect mesh-covered holes before exiting into the main recirculating system. Sea hares of different size classes (< 100, 100—200, 200-300, 300-400, > 400 mg ww, n = 35 sea hares) were placed into contact with pycnogonids at the start of each trial. Experimental compartments were checked over the fol- lowing 2 days, and the condition of each sea hare re- corded. Chemical Effects To determine if the presence of different types of algal secondary metabolites (i.e., not from Laurencia obtusa) deterred Anoplodactylus evansi from feeding, pycnogo- nids were offered Aplysia parvula that had fed on the red seaweed Delisea pulchra. D. pulchra contains halogenat- ed secondary metabolites, and A. parvula accumulates high concentrations of these compounds in its tissues (de Nys et al., 1996), significantly more so than Laurencia obtusa metabolites (Rogers et al., 2000). D. pulchra did not occur at the site where A. evansi was found, nor in that area of Port Jackson, so the secondary metabolites from this seaweed may be regarded as novel to the pyc- nogonids used. However, A. evansi would undoubtedly encounter D. pulchra in its wider geographic range, and consequently, pycnogonids from the study area in Port Jackson may possibly have an innate response to D. pul- chra metabolites. Because we cannot rule out such a pos- sibility, the term “novel” in reference to D. pulchra should be interpreted at the least as meaning it did not co-occur with the pycnogonids tested. Five individual pycnogonids were maintained in separate beakers and of- fered five replicate juvenile A. parvula (100—200 mg ww) collected from D. pulchra at Bare Island. Each sea hare was placed in contact with a pycnogonid, and the result of the trial recorded after 2 days. To determine if increased quantities of compounds in (<\South Head Port Jackson 33°51'S Shark Bay|/ A B Shark Island D 151 16E |S Figure 1 The position of study sites A, B, C, and D near the entrance to Port Jackson, east of Sydney, New South Wales, Australia. sea hares deterred feeding by Anoplodactylus evansi, pyc- nogonids were held in individual cells in 25 L experi- mental aquaria and fed sea hares according to the follow- ing experimental design. Pycnogonids were offered either sea hares treated with adult extracts (1.e., to double the level of extract present), untreated sea hares (1.e., average The Veliger, Vol. 43, No. 4 levels), or solvent-treated sea hares as a control. A further group of solvent-treated sea hares was held separately in aquaria cells without pycnogonids to control for “‘auto- genic”? changes in the weight of sea hares unrelated to consumption by pycnogonids. Ten replicate animals were given each treatment. The Aplysia parvula used in this experiment were juveniles collected from Delisea pulchra (mean 123 + SE 15 mg ww, which contain on average 4.3 + SE 0.8 mg of non-polar extract). The juvenile sea hares were frozen, then thawed prior to use. The non- polar extract used to treat sea hares was obtained from adult A. parvula from D. pulchra by sequential extraction of freeze-dried animals, using 3 xX 10 mL aliquots of dichloromethane (AR). The resulting solutions were com- bined, syringe filtered (PTFE 0.5 pm), then air dried. Crude extracts from sea hares contained on average 42.2 + SE 4.5% total secondary metabolites, and juvenile and adult sea hares had similar levels of secondary metabo- lites in their extracts. The relationship between crude ex- tract levels (y in mg) and sea hare wet weight (x in g) was determined for D. pulchra fed sea hares by linear regression (y = 39.9 X +0.1, R* = 0.86, p < 0.001, n = 42). To double the quantity of extract per sea hare in the extract treatment group, the quantity of crude extract pre- sent in each juvenile sea hare was calculated, then an equivalent amount of adult extract was injected into the body cavity of the sea hare. Ethanol was the solvent used to apply the extract, and was also injected into the solvent control and autogenic control treatments at equivalent volumes to those used to administer the extract. The weight of sea hares was measured at the start of the ex- periment and after 2 days, and the proportional change in weight calculated. Table 1 Opisthobranch mollusks consumed by the pycnogonid Anoplodactylus evansi, grouped by order and family. Published maximum lengths for each species are included (After Burn, 1989; Coleman, 1989; Wells & Bryce, 1993). Maximum length Order—Family Species (mm) Anaspidea Aplysiidae Aplysia parvula Guilding in Morch, 1863 60 Aplysia dactylomela Rang, 1828 250 Aplysia sydneyensis Sowerby, 1869 150 Aplysia juliana Quoy & Gaimard, 1825 200 Stylocheilus longicauda Quoy & Gaimard, 1824 40 Ascoglossa Oxynoidae Oxynoe viridis Pease, 1861 10 Elysiidae Elysia australis Quoy & Gaimard, 1832 10 Nudibranchia Polyceridae Plocamopherus imperialis Angas, 1864 100 Chromodorididae Hypselodoris bennetti Angas, 1864 50 Glossodoris atromarginata Cuvier, 1804 50 Bornellidae Bornella stellifer Adams, 1848 40 Glaucidae Austraeolis ornata Angas, 1864 35 Aeolidiidae Spurilla australis Rudman, 1982 55 C. N. Rogers et al., 2000 To examine whether algal secondary metabolites are sequestered by pycnogonids after feeding, chemical anal- yses were done on pycnogonids that had consumed a sea hare and compared to analyses of starved pycnogonids. The Anoplodactylus evansi used were maintained aerated in 2 L beakers with a piece of the red alga Laurencia obtusa as habitat. Each treatment consisted of Aplysia parvula or Aplysia dactylomela Rang, 1828, (both col- lected from L. obtusa) or starved controls. Three repli- cates were given each treatment. The pycnogonids were allowed to consume the sea hares over 2 days, then were frozen for extraction. The pycnogonids were freeze dried, crushed, then sequentially extracted using 3 X 5 mL al- iquots of dichloromethane (AR). The resulting extract was syringe filtered, then air dried and weighed. Samples were prepared for gas chromatography mass spectrometry (GCMS) by dissolving the extract in analytical grade eth- yl acetate (0.2 mg/mL) containing 10 g/mL naphthalene internal standard. The samples were then analyzed for Laurencia obtusa secondary metabolites using a Hewlett Packard HP5980 series II gas chromatograph and HP5972 mass selective detector, following the procedures of de Nys et al. (1998). The Abundance of Anoplodactylus evansi and Aplysia parvula on Laurencia obtusa The alga Laurencia obtusa was collected to measure the abundance of Anoplodactylus evansi and Aplysia par- vula during consecutive years from three sites (A—C in Figure 1) in Port Jackson, NSW. Laurencia obtusa was chosen to assess the abundance of both A. evansi and A. parvula because both species were common on this alga. Algal thalli were removed from the substratum, carefully placed into individual plastic bags with seawater, and then taken to the laboratory for sorting. The number of pyc- nogonids and sea hares present on each thallus was re- corded, and the wet weight of each thallus was measured. Sampling was commenced at site C at Parsley Bay when no L. obtusa were found at site B during 1996. Statistical Analyses Results of predation trials on different sizes of Aplysia parvula by Anoplodactylus evansi were compared in a contingency table for sea hares larger and smaller than 200 mg ww. A G-test was used to test the hypothesis that consumption of sea hares by A. evansi was equivalent for both size groups. Pycnogonid consumption of extract-treated versus oth- er Aplysia parvula was done using a single factor analysis of variance (ANOVA) followed by Tukey’s HSD test at the a = 0.05 significance level. Data for the proportional change in weight of sea hares were transformed using Ln(1 + x) before analysis and checked for homogeneity of variance using Cochran’s test. The initial size of sea hares in each treatment was compared using a single fac- Page 333 tor ANOVA, and was not significantly different between treatments (F; 3, = 2.14, P = 0.113). Field data for the size of Laurencia obtusa thalli, and abundance of Aplysia parvula were analyzed using a sin- gle factor ANOVA comparing the sites in Port Jackson, followed by Tukey’s HSD test. Data for abundance of Aplysia parvula were Ln(1 + x) transformed. Because sites were not sampled for all dates, sampling dates were pooled for each site in comparisons. Data for abundance of A. evansi were not homogeneous using Cochran’s test because Anoplodactylus evansi only occurred at one site, sO a non-parametric test (Mann-Whitney) was used to compare abundance of pycnogonids between site A ver- sus sites B and C. RESULTS The pycnogonid Anoplodactylus evansi was found only at site A in Shark Bay, Port Jackson, close to Shark Island (Figure 1) where the type specimens were collected (Clark 1963). This species was not observed at nearby sites B, C, or D (Figure 1), or at other localities in Botany Bay and Port Hacking, south of Port Jackson. The pyc- nogonids were found inhabiting several species of algae, including the brown algae Sargassum linearifolium Turn- er, 1809, Sargassum vestitum R. Brown, 1811, Padina crassa Yamada, 1931, Dictyopteris acrostichoides J. Agardh, 1882, as well as the red alga Laurencia obtusa and an arborescent bryozoan. Pycnogonids were observed to be nocturnally active, as were juvenile Aplysia parvula and Aplysia dactylomela. During the day, pycnogonids and sea hares were found sheltering in the basal sections of algae. Anoplodactylus evansi consumed 13 different species of opisthobranch mollusk from eight families, encom- passing three orders (Table 1). Of the species eaten, Aply- sia parvula and the nudibranch Bornella stellifer Adams, 1848, were collected most often at site A where A. evansi occurred. Anoplodactylus evansi was also observed to consume an unidentified errant polychaete worm. Pyc- nogonids did not consume small (5—15 mm in length) amphipods, prosobranch mollusks, or echinoids that were offered to them. A. evansi consumed only small individ- uals of each opisthobranch species. Those eaten were generally less than 200 mg ww (< 10 mm in length), which for most species was much smaller than adult spec- imens observed in the field or published maximum sizes (Table 1) (Burn, 1989; Coleman, 1989; Wells & Bryce, 1993). When A. evansi was offered A. parvula, ranging in size from 20 to 600 mg ww, the pycnogonids could subdue and consume only animals smaller than 300 mg ww (Figure 2). The effect of prey size (< 200 versus > 200 mg ww) on consumption by A. evansi was significant (G = 8.53, P < 0.005). At adult size this pycnogonid species has a leg span of up to 30 mm across and weighs Page 334 % eaten <100 100-200 200-300 300-400 > 400 Sea hare live weight (mg) Figure 2 Consumption of different sizes of the sea hare Aplysia parvula by Anoplodactylus evansi. Bars show percentage of individuals eaten; the number of sea hares per sample is shown in brackets above the bar, (n). 50-80 mg ww. A. evansi feeding on 300 mg sea hares is attacking prey five to six times its own body weight. Anoplodactylus evansi subdues prey using the front four legs to grasp and hold, while the back four legs remain secured to the substratum. Each leg ends with a long claw, which can be clamped against an opposing clawed palm, giving the pycnogonid a secure hold on slimy opisthobranchs. Once a prey item is held, A. evansi uses its chelicerae to tear pieces of flesh from the prey, which are passed to the proboscis, or alternatively, the proboscis is applied directly to the prey. While A. evansi typically consumed all of the soft tissue, some sea hare digestive glands were not consumed (33% of cases, n = 18). Sea hares often released ink when attacked, although the crouching pycnogonid was generally well clear of any secretions. Sea hare secretions did sometimes contact a front leg, and pycnogonids responded to such contact by vigorously waving the affected limb, as though irritated. Where pycnogonids were held together in the same hold- ing tank it was noted that they would feed in groups on a prey item. In the trial to determine if sea hares that had fed on a “novel” algae (Delisea pulchra) would be consumed, Anoplodactylus evansi ate all five juvenile Aplysia par- vula from D. pulchra that were offered. However, A. evansi was deterred from feeding on A. parvula that had artificially increased levels of secondary metabolites (Fig- ure 3). Pycnogonid feeding was significantly less on sea hares that had double extract levels compared to either solvent-treated or untreated sea hares (Figure 3). The sea hares treated with adult extract had similar weight loss to the autogenic control animals which were kept separately, without pycnogonids indicating that little, if any, feeding occurred. Pycnogonids which had fed on Aplysia parvula or A. The Veliger, Vol. 43, No. 4 S ‘p Weight loss (prop.) So Ww 0.2 0.1 0 Auto-control 2x extract Solvent Untreated Treatment Figure 3 Mean weight loss (proportion) from Aplysia parvula + 1SE, for treatments offered to Anoplodactylus evansi (except autogenic controls). The result of a one way ANOVA on Ln(1 + x) trans- formed data was F;,, = 7.94, P < 0.01. Treatments which are not significantly different in weight loss at a = 0.05 using Tu- key’s test share the same letters on the graph. n = 10 per treat- ment. dactylomela containing Laurencia obtusa secondary me- tabolites contained trace amounts of the terpene palisadin A at levels below the quantitative limit of the GCMS. Such small quantities of this secondary metabolite could be attributed to sea hare tissue in the pycnogonids’ gut or to low levels of accumulation. Pycnogonids which were starved as controls contained no detectable peaks for L. obtusa secondary metabolites on chromatographs of samples. Collections of the red alga Laurencia obtusa at three sites (A—C in Figure 1) during late summer/autumn from 1995 to 1998 had significantly lower abundances of the sea hare Aplysia parvula on thalli where Anoplodactylus evansi occurred (Figure 4, Table 2). There was no sig- nificant difference between sites in the size of thalli col- lected (Figure 4a, Table 2). L. obtusa thalli at site A were observed to persist during the winter, whereas thalli at the other sites disappeared. A. evansi occurred only at site A. The abundance data for A. evansi were not homogeneous using Cochran’s test, so a Mann-Whitney test with normal approximation was used to compare the abundance of pycnogonids between sites (A vs. B and C). The result was a significant difference in pycnogonid abundance be- tween site A versus sites B and C (Z = 6.23, P < 0.001). DISCUSSION Anoplodactylus evansi is a generalist predator of small opisthobranchs, and other soft-bodied invertebrates, as are other Anoplodactylus spp. (King, 1973; Piel, 1991). This pycnogonid hunts opisthobranchs on benthic algae, immobilizing them with movable claws on the front legs, before consuming them. A. evansi consumed whole opis- C. N. Rogers et al., 2000 Page 335 (A) 30 25 Laurencia obtusa 20 15 10 Plant wet weight (g) A-5/95 A-5/96 A-6/97 A-4/98 B-5/95 B-4/98 C-5/96 C-2/97 C-4/98 Aplysia parvula Abundance (number / g) A-5/95 A-5/96 A-6/97 A-4/98 B-5/95 B-4/98 C-S5/96 C-2/97 C-4/98 Anoplodactylus evansi (C) 0.8 i S fon oS ty Abundance (number / g) So ay A-5/95 A-5/96 A-6/97 A-4/98 B-5/95 B-4/98 C-5/96 C-2/97 C-4/98 Sample site-date Figure 4 (A) Mean size of Laurencia obtusa (g ww) + ISE at sampling sites in Port Jackson. Samples identified by site letter and sample date (month/year). Number of plants per sample is shown in brackets above bars, (n). (B) Abun- dance of Aplysia parvula on Laurencia obtusa (number of sea hares/g ww) + 1SE at sampling sites in Port Jackson. (C) Abundance of Anoplodactylus evansi on Laurencia obtusa (number of pycnogonids/g ww) + 1SE at sampling sites in Port Jackson. Page 336 Table 2 Analysis of variance comparing: (A) thalli size of Lau- rencia obtusa at sites in Port Jackson, and (B) the abun- dance of Aplysia parvula on L. obtusa (Ln(x + 1) trans- formed) at sites in Port Jackson. Sampling dates were pooled for each site in analyses. The results of a Tukey’s HSD test showed the abundance of Aplysia parvula at site A was significantly different (a = 0.05) from sites B and C. Species 4 (A) L. obtusa (B) A. parvula Source of variation df F P df 1 P Between groups 2 0.85 0.429 2 13.44 < 0.001 Within groups 94 94 thobranchs, including the nudibranchs Bornella stellifer, Austraeolis ornata Angas, 1864, and Spurilla australis Rudman, 1982, not just the cerata as was observed for Anoplodactylus carvalhoi (Piel, 1991). The digestive gland of some sea hares was not eaten by A. evansi. Aply- sia parvula stores the bulk of acquired algal secondary metabolites in its digestive gland (de Nys et al., 1996), so pycnogonids would avoid exposure to high concentra- tions by rejecting this organ. Only small opisthobranchs less than 300 mg were eaten, as larger animals could not be subdued, indicating an effect of prey body size. Pen- nings (1990a) also found an effect of prey body size on the capture success of opisthobranchs by Aglaja inermis Cooper, although in this instance, smaller individuals were consumed less often because habitat complexity aid- ed predator evasion. Group feeding by A. evansi may fa- cilitate (by weight of numbers) successful attacks on opis- thobranchs larger than the prey size range found here. The nocturnal vertical movement of A. parvula (Rogers et al., 1998) and nocturnal activity of A. evansi may also increase the encounter rate between predator and prey in this system. The local distribution of Anoplodactylus evansi was re- stricted, with pycnogonids found only at one of four near- by sites that had similar benthic communities (Figure 1). A. evansi has been collected at several sites along the New South Wales coastline, and as far south as Tasmania (Clark, 1963), so its limited distribution in Port Jackson contrasts with its wide distribution along the adjacent coastline. The limited distribution of this pycnogonid is most probably due to its brooding reproduction (King, 1973), and consequent limited dispersal to nearby sea- weed. At site A, where A. evansi was found, pycnogonids occurred on several different seaweeds and a bryozoan, indicating that a broad range of organisms is used as hab- itat. Natural levels of secondary metabolites and other diet- The Veliger, Vol. 43, No. 4 derived defenses of the opisthobranch mollusks investi- gated here had no effect in deterring Anoplodactylus evansi from attacking and consuming them. The small opisthobranchs consumed by A. evansi may lack suffi- cient quantities of such compounds to form an effective deterrent, although some juvenile sea hares can have high concentrations of secondary metabolites (Rogers, unpub- lished data). Artificially increasing the levels of metabo- lites present to double their average quantity did deter A. evansi from feeding on juvenile Aplysia parvula, dem- onstrating a quantitative deterrent effect. A possible rea- son why A. evansi did not consume all sea hares (< 300 mg ww) offered to them (see Figure 2) may be because the rejected sea hares had higher levels of secondary me- tabolites. Hence, concentration of metabolites can reduce feeding by A. evansi, but most A. parvula outgrow the prey range of this pycnogonid before they can store the levels of metabolites necessary for deterrence. A similar process may occur for the other opisthobranchs investi- gated, except for the ascoglossans, which attain much smaller maximum sizes (Table 1). The abundance of Aplysia parvula was significantly lower on the red alga Laurencia obtusa at site A over consecutive years compared to other nearby sites where Anoplodactylus evansi did not occur. L. obtusa thalli from site A were similar in size to thalli at the other sites sam- pled. L. obtusa thalli inhabited by A. evansi were ob- served to persist into the winter, whereas thalli at other sites disappeared. The disjunct distribution of the pyc- nogonid A. evansi and sea hare A. parvula is consistent with this predator excluding A. parvula from site A, al- though other explanations are possible. Anoplodactylus evansi is able to tolerate low levels of secondary metabolites and feed successfully on a wide variety of small opisthobranchs with different defensive substances. The opisthobranchs investigated here are known to sequester different metabolites (Gunthorpe & Cameron, 1987; Avila, 1995; de Nys et al., 1996), and the juvenile Aplysia parvula tested here contained sec- ondary metabolites from Laurencia obtusa or Delisea pulchra, but all were consumed nonetheless. The pres- ence of “novel”? secondary metabolites from D. pulchra, which did not co-occur with A. evansi in Port Jackson, suggests that this pycnogonid can tolerate a variety of secondary metabolites in its diet. Given the wide range of opisthobranchs eaten by A. evansi in this study, such compounds may also include those found in sponges, cni- darians, bryozoans, and other dietary items of the opis- thobranchs. One reason this pycnogonid can feed suc- cessfully on potentially poisonous opisthobranchs could be their digestive processes involving pinocytosis and in- tracellular digestion of food particles (Richards & Fry, 1978). Isolation of prey tissue containing secondary me- tabolites in vacuoles by pycnogonids, with eventual ex- cretion of the residual body and presumably any toxins, would render such substances harmless. Disposal of sec- C. N. Rogers et al., 2000 Page 337 ondary metabolites by this digestive process may also account for the trace amounts of compounds found in A. evansi which had consumed Aplysia. The pycnogonid A. evansi is thus a generalist predator of opisthobranchs, which may overcome their defenses by virtue of an iso- lating digestive process and by consuming small individ- uals which may lack substantial quantities of diet-derived defenses. Acknowledgments. We thank Alistair Poore for his encourage- ment to investigate Anoplodactylus, Timothy Charlton for pro- cessing GCMS samples, Dr W. Rudman for help with identifi- cation of opisthobranchs, and two anonymous reviewers whose comments improved this manuscript. C. N. Rogers was supported by an Australian Postgraduate Award during the period of this work. R. de Nys was supported by an Australian Research Coun- cil Research Fellowship. This study was funded by Australian Research Council grant # A19530672 to P. D. Steinberg. LITERATURE CITED AvILA, C. 1995. Natural products of opisthobranch molluscs: a biological review. Oceanography and Marine Biology An- nual Review 33:487—-559. Burn, R. D. 1989. Opisthobranchs. 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D. 1991. Diet specialization limits herbivorous sea slugs’ capacity to switch among food species. Ecology 72:1880-1888. TROWBRIDGE, C. D. 1994. Defensive responses and palatability of specialist herbivores: predation on NE Pacific ascoglossan gastropods. Marine Ecology Progress Series 105:61—70. VAN DER VELDE, J. T. & R. J. Kinc. 1984. The subtidal seaweed communities of Bare Island, Botany Bay. Wetlands 4:7—22. WELLS, E E. & C. W. Bryce. 1993. Sea Slugs of Western Aus- tralia. Western Australian Museum: Perth. 184 pp. THE VELIGER © CMS, Inc., 2000 The Veliger 43(4):338-348 (October 2, 2000) Life History of a Hydroid/Nudibranch Association: A Discrete-Event Simulation CHARLES M. CHESTER*, ROY TURNER’, MICHAEL CARLE!' anp LARRY G. HARRIS! ‘Department of Zoology, University of New Hampshire, Durham, New Hampshire 03824, USA "Department of Computer Science, University of Maine, Orono, Maine 04469, USA Abstract. The general paradigm for early community succession is that early colonists do not replace themselves. Hydroids are typical early colonists that play an important role in the recruitment of later species. Aeolid nudibranchs are important predators on hydroids, and may thus have an indirect impact on succession. In an attempt to understand the mechanics of community change, we modeled nudibranch-hydroid community dynamics using a discrete-event simulation. Data for the model was obtained from life history studies of the aeolid nudibranch Tenellia adspersa (Nord- mann, 1845) and its hydroid prey Cordylophora lacustris (Allman, 1853). Seven simulations were performed, varying adult immigration, emigration, and larval settlement. The results of these simulations have important implications for early community succession and the role of nudibranchs in the persistence of hydroid colonies. In all simulations, the hydroid colony was completely removed by T. adspersa. Cordylophora lacustris persisted for the longest time in simulations with no adult migration or larval settlement. In addition, nudi- branchs persisted for up to 46 days after their food supply was exhausted. These predictions suggest that Tenellia adspersa can play an important indirect role in succession by removing hydroids and preventing their re-establishment. INTRODUCTION The general paradigm for early community succession is that early colonists do not replace themselves (Connell, 1975). Connell (1978) suggested that succession could vary depending on factors such as disturbance, competi- tive interactions, and temporal availability of propagules. Connell & Slatyer (1977) proposed that established spe- cies could inhibit, facilitate, or remain neutral in the re- cruitment of later successional stage species. Hydroids are typical early colonists in fouling com- munities; they tend to be succeeded by other species such as barnacles, tunicates, and mussels (Harris & Irons, 1982). Hydroids often have ephemeral life-history and distribution patterns, but can be important in affecting the recruitment of later successional species (Standing, 1976; Chester, 1996a). The majority of aeolid nudibranchs are partial preda- tors, consuming portions of hydroid colonies (Todd, 1981; Todd, 1983). Aeolid nudibranchs play a significant role in structuring hydroid communities (Harris, 1987). Aeolid predation can create physical gaps in prey colo- nies, alter the population structure, or cause changes in the prey’s growth form (Gaulin et al., 1986). The impact of nudibranchs on hydroid colonies varies in relation to the number of predators in the colony. In low numbers, * Present Address: Department of Biology, Spring Hill Col- lege, Mobile, Alabama 36608, USA. the impact may be limited because the hydroid colony can grow faster than the nudibranch predation rate. How- ever, at higher abundance, the impact is more substantial and will ultimately lead to the removal of the hydroid colony. A number of aeolid species have life histories with short generation times and high reproductive output, which enable them to take advantage of temporally un- predictable, but often abundant, food resources. Most of these species have an obligate planktotrophic larval stage capable of remaining in the plankton for weeks to months. At least one of them, Tenellia adspersa (Nord- mann, 1845), has lecithotrophic larvae that are capable of metamorphosing within the egg capsule (Chester, 1996b). This has important implications for nudibranch popula- tion growth within a hydroid colony and for the persis- tence of that colony. For populations of aeolids having pelagic larval stages, population growth in a hydroid col- ony will be determined by settlement and metamorphosis of larvae from the plankton. For aeolids having capsular development (or non-pelagic lecithotrophic develop- ment), population growth following initial recruitment will be determined by growth and reproductive rates of resident individuals within a hydroid colony. In the latter case, the aeolid’s populations will increase at an expo- nential rate, inundating the hydroid colony and consum- ing it in a relatively short time. Tenellia adspersa is a small (5-7 mm) aeolid nudi- branch commonly found in New England estuarine en- C. M. Chester et al., 2000 vironments (Clark, 1975). Tenellia adspersa is a gener- alist that feeds on a variety of gymnoblastic and calyp- toblastic hydroids. Most of these hydroids are seasonally abundant in the Great Bay Estuary, New Hampshire (70°55'N, 43°5'W), existing as colonies on piers, floating docks, eelgrass blades, and other natural and artificial structures. In the estuary, the distribution and abundance of these hydroids can change within as little as one week’s time (Chester, 1996a). Tenellia adspersa has a plastic developmental mode where both pelagic lecitho- trophic larvae and benthic juveniles are produced in the same spawn mass and by the same individual (Eyster, 1979; Chester, 1996b). This study explores the implications of nudibranch life history on the persistence of a hydroid colony. The goal of this study was to design a computer simulation that would model hydroid-nudibranch dynamics. The study involves both laboratory studies and computer models of the nudibranch-hydroid system. MATERIALS anp METHODS The hydroid, Cordylophora lacustris (Allman, 1853), was collected in September 1991 from a floating dock on the Oyster River, Durham, New Hampshire (70°55’N, 43°08’W). Colonies were cultured on glass slides that were suspended in aquaria at 25°C and at a salinity of 15-25%o. A small portion of a colony (~ 3 polyps) was placed under a monofilament line that was tightly wrapped around the slide. Colonies were fed nauplii of Artemia salina every other day. The nudibranch Tenellia adspersa was collected in Oc- tober 1991 from within C. lacustris colonies on a floating dock on the Squamscott River, Rockingham, New Hamp- shire (70°56'N, 43°02’W). Stock cultures were estab- lished in fingerbowls at 25°C and 25%o and provided with an ad libitum amount of C. lacustris. For the computer simulation comparing nudibranch and hydroid dynamics, the following observations were gathered: (1) measurement of hydroid colony growth, (2) determination of nudibranch life history, and (3) mea- surement of predation rates on hydroid colonies. These data were used to construct a discrete-event simulation model (Banks et al., 1996) of nudibranch population dy- namics within a single hydroid colony. This type of mod- el deals with individuals rather than aggregate behavior, thus it provides a finer-grained simulation than typical analytical models. It treats time as discrete units, with events determining its passage; thus, it simplifies the model enough to be tractable. Discrete-event simulation is often performed when the system being simulated is very complex, when some of the information needed for an analytical model is missing, or when it is important to track the effects of individual differences in simulated entities. Page 339 Hydroid Colony Growth Small pieces of C. lacustris containing approximately three polyps were attached to glass slides and suspended in aquaria at 20 and 25%o, with 10 replicates per salinity. These colonies were fed A. salina nauplii every other day. The number of polyps was counted every few days, and each colony was mapped at 60 using a Wild dissecting microscope equipped with a drawing tube. Stolon length was measured with a map measurer, and the distance con- verted to the nearest 0.1 mm. In C. lacustris, stolons and uprights grow at a fairly constant rate (Fulton, 1963). Therefore, the inside diameter of stolon was measured to the nearest 1 wm from histological cross-sections with a compound microscope equipped with an ocular micro- meter (diameter = 294 + 2 wm, n = 10), and this value was used to calculate volume of stolon. Nudibranch Life History Five newly laid spawn masses from Tenellia adspersa were haphazardly collected from the stock culture and raised with an ad libitum amount of C. lacustris in the same conditions as the stock culture. Water was changed daily and the number of nudibranchs recorded. Daily ob- servations were made on nudibranch body length, number of spawn laid and number of eggs per spawn with a bin- ocular dissecting microscope equipped with an ocular mi- crometer. Nudibranch Feeding Rates In order to measure feeding rates, nudibranchs of var- ious sizes were placed in colonies of C. lacustris and followed over time. The number of polyps was counted daily and the amount of living stolon measured as with the hydroid colony growth. The feeding rate was calcu- lated as the change in living stolon, and represents the amount of growth less the amount eaten by the nudi- branch. Discrete-Event Simulation Model A discrete-event simulation model (e.g., Banks et al., 1996) was constructed to model nudibranch population dynamics. In the current case, the individuals modeled were nudibranchs. The information modeled per nudi- branch included: time of birth, time of metamorphosis, number of spawn (egg masses), current reproductive rate (spawn/day, eggs/spawn), starvation status (including du- ration of starvation), and reproductive shut-off due to starvation. These parameters were changed during the simulations depending on the amount of food available to each nudibranch. As our primary goal was to model the effect of indi- vidual nudibranchs on a hydroid colony, the hydroid col- ony was treated as a bulk food source. The amount of the hydroid was updated at the end of each simulated day Page 340 The Veliger, Vol. 43, No. 4 Table 1 Parameters used in the discrete-event simulation experiments. Values were obtained from the laboratory growth and feeding experiments. Nudibranchs fed Parameter Mean Adult lifespan (days) 24.5 Time to metamorphosis (days) deal Time to first spawn (days) 17.6 Number of spawn/day 5.4 Eggs per spawn 32.1 Hatching time (days) 6.3 Day of first feeding 5.0 Percent of spawn that hatch 100.0 Percent pelagic lecithotrophic larvae 76.8 Hydroid growth rate (%/day) 22. Feeding rate (mm hydroid/day * nudibranch) by nudibranch size 0-1 mm 0.3 1-2 mm 4.7 2-3 mm 4.9 3-4 mm 7.4 >4mm 18.0 based on the hydroid’s growth that day and the total amount eaten by all nudibranchs. The parameters used in the simulations were based on SALE.: the laboratory experiments and are shown in Table 1. All parameters were held constant across simulation experi- ments except the independent variables: immigration rate 41), emigration rate (E), and larval settlement rate (S). Each simulation started with two adult nudibranchs and was allowed to run either for 80 days or until there were no events in the simulator’s event queue other than SLE, ‘“‘*housekeeping”’ events (such as data collection). This corresponded to the situation of there being no nudi- branchs of any life stage other than pelagic veligers in the system. The dependent variables of interest were N.,,, SDE. : the number of nudibranchs remaining alive at the end of the experiment, and t,, the time between food source ex- tinction and nudibranch population extinction (in the cas- es where N.,, = 0). Seven simulation experiments were performed as follows: SAE. : S.[.E,: No immigration or emigration of adults, no larval settlement (in all experiments, hatched veligers were assumed to enter the plankton). This is similar to the laboratory experiments. No immigration or emigration of adults, but larval settlement at a rate of one to three veligers per day (2.0 + 0.1), which is con- sistent with what was observed in the Great Bay Estuary during the peak summer months (Chester, personal observation). No immigration or emigration when food was present; emigration (10% per day) when SLES (5 oped 9 Saabs: Nudibranchs starved Std. Err. 1.0 0.2 0.3 0.9 0.6 0.2 Mean Std. En. 0.3 25g 4.8 0.3 lea/ 0.3 1.4 no food is available; and larval settlement at the rate above with or without food. Same as S,I,E,, except that larval settlement occurred only when food was present. This might correspond, for example, to an isolat- ed population of nudibranchs where no other populations are close enough to allow im- migration, but where veligers are present in the water column. : Immigration of 2.0 + 0.1 adults per day, ran- dom ages; a constant, low emigration rate (1% per day); and larval settlement. All oc- curred regardless of food source status. Immigration at the above rate only when food is present; low emigration rate (1% per day) when food is present and a higher rate (10% per day) when food is exhausted; and larval settlement at the rate above regardless of food status. Same as simulation S.I,E.,, except that larval settlement occurs only when food is present. This is conjectured to be similar to the case in the field. Simulations were run both using the means of all pa- rameters (the ‘“‘means-only”’ case) and using the standard errors to determine normal distributions of the parameters (the ‘‘normal-distribution”’ case). It should be noted that even in the means-only runs, there is still some variation due to the means of integer parameters (e.g., eggs/spawn) being real numbers. In these cases, a probabilistic round- ing scheme was used. Fifty simulations were run for each C. M. Chester et al., 2000 case of each experiment, and the results averaged. Initial food supply in the means-only case was sufficient for 15 nudibranchs for one day, whereas in the normal-distri- bution case it was roughly half that, sufficient for seven nudibranchs for one day. This was because with the larger amount of food, in the normal-distribution case occasion- ally the population would become too large to simulate practically on the available computers. Statistical Analysis Statistical analyses of the laboratory studies were per- formed using SYSTAT (vers. 5.03, Systat Inc., Evanston, Illinois). The relationship between nudibranch size and predation rates was investigated with an analysis of var- iance model (ANOVA) (Sokal & Rohlf, 1981; Zar, 1996). Tukey HSD was used to compare nudibranch sizes with different feeding rates. The discrete-event simulation re- sults were analyzed using CLASP (Common Lisp Ana- lytical Statistics Package) (Anderson et al., 1994). Means and standard errors are used throughout. RESULTS Tenellia adspersa is a small nudibranch, reaching a max- imum size of 6-8 mm. The body bears five to seven clus- ters of cerata, usually with two to three cerata per cluster. Cnidosacs are present at the tips of each cerata. The tip of the penis is armed and bears a cuticular stylet. Hydroid Colony Growth The most rapid growth in Cordylophora lacustris oc- curred at 20%c salinity (Figure 1A, B). Higher increase in the number of polyps occurred at 20%c, with slower growth at 25%c (Figure 1A). This corresponds to an 18.1 * 0.5 percent increase at 20%c and a 4.5 + 0.4 percent increase at 25%c. Stolon growth closely followed growth of polyps, with the more rapid growth occurring at 20%o (Figure 1B). At these salinities, stolons grew 2—3 mm/?/ day. Slower growth occurred at 25%c (stolon growth: 0.5 + 0.3 mm?/day). The percent change in volume of stolon was 21.2 + 1.4 at 20%o and 7.2 + 0.4 at 25%o. Nudibranch Life History The results of the life history study are presented in Figure 2. The generation time from egg to egg was ap- proximately 17.6 + 0.3 days (n = 20) and the life cycle from egg to death was 24.5 + 1.0 days (n = 10). Hatch- ing occurred in 6.3 + 0.2 days (n = 56). Larvae meta- morphosed in 7.1 + 0.2 days (n = 45) at a juvenile size of 0.3 + 0.1 mm (n = 20). Tenellia adspersa grew ex- ponentially in size until sexual maturity, as observed by the presence of the first spawn mass. Size at maturity was 4.5 + 0.2 mm (n = 20). The growth rate decreased after sexual maturity until a maximum length of 6.1 + 0.3 (n = 10) was achieved. Nudibranchs decreased in size near Page 341 # of Polyps Volume of Stolon (mm*) 0 5 10 15 20 25 Time (days) Figure 1 Growth of Cordylophora lacustris at two salinities, measured as number of polyps (A), and volume of stolon (B). the end of their life, reaching a length of 5.4 + 0.3 mm (n = 10). This decrease became apparent up to 7 days before death. Mature nudibranchs produced 5.4 + 0.9 spawn masses per day (n = 10) with 32.1 + 0.6 eggs per spawn (n = 10). Their lifetime fecundity was 39.4 + 7.6 spawn per individual (n = 10) for a total of 1269.3 + 102.9 eggs per individual (n = 10). Page 342 The Veliger, Vol. 43, No. 4 Nudibranch Feeding Rate The results of the predation experiments are presented in Table 1 and Figure 3. Tenellia adspersa feeds by pierc- ing the perisarc of the stolon with its radula and sucking out tissue. Larger nudibranchs (> 6 mm) were observed to feed on the polyps themselves by rasping bites out of the polyp. Newly metamorphosed juveniles were invari- ably found near new hydroid growth. No observable loss of hydroid tissue was observed for the first 4—5 days fol- lowing metamorphosis. Nudibranchs ate significantly more polyps and more coenosarc as they grew (ANOVA: polyp predation, F,,, = 168.0, P < 0.0001; stolon pre- dation, F,,, = 22.9, P < 0.0001) (Figure 3). Mature nu- dibranchs (> 4 mm) consumed 18.0 + 1.9 mm: of stolon tissue per day and 6.6 + 0.2 polyps per day. Discrete-Event Simulation Experiments Figures 4 and 5 illustrate the output of the simulator for each experiment. The results, including the maximum nudibranch population sizes, are summarized in Table 2. Each experiment is discussed briefly below: Experiment S,I,E,. In the means-only case (Figure 4A), the nudibranch population went extinct long after the food supply was exhausted (t, = 24.4 + 0.3 days). This difference between the two extinction times was signifi- cant (two-tailed t-test, P < 0.0001). The population re- mained stable after the last spawn had hatched until about 45 days, then dropped rapidly to zero. In the normal dis- tribution case (Figure 4B), there were remnants of the population that remained alive in some of the runs, giving Ning = 0.5 + 0.5. There appears to be two generations of nudibranchs as opposed to one in the means-only case as observed by two peaks in spawn masses. In this case as well, the nudibranch population outlived the hydroid by a large, significant time (t, = 45.9 + 0.8, P < 0.0001). The fact that t, in this case is greater than the mean li- fespan of the nudibranchs is due in part to the presence of spawn that hatched following food exhaustion. Experiment S.J,E,. The graph of the means-only case was very similar to that of experiment S,I,E,, except that there was still a small, nearly constant nudibranch pop- ulation (N.,,4 = 34) present at the end of the experiment due to a constant influx of larvae from the plankton. This is also true of the normal-distribution case (N.,4 = 44.0 += (0:6): Experiment S.I,E,. In the means-only case (Figure 4C), the population decreased smoothly until about day 43, at which point there is a fast decline. This is most likely due to nudibranchs beginning to die off and emigrate. There is a small, nearly constant population of nudi- branchs (N.,4 = 14.9 + 0.2) left at the end of the exper- iment. The normal-distribution case is similar, with N.,, = 20.5 + 0.4 (Figure 4D). Experiment S,I,E,. Both the means-only (Figure 4E) and the normal-distribution cases (Figure 4F) were very similar to their counterparts in S.I,E,, except that the nu- dibranchs became extinct in both cases long after the food was exhausted (ty = 24.9 + 0.1 and 30.7 + 0.6, respec- tively, P < 0.0001). Experiment S.J.E.. Both the means-only (Figure 5A) and the normal distribution case (Figure 5B) show vir- tually no decrease in nudibranchs after the final build-up of the population once the food is exhausted. Immigra- tion, emigration, and larval settling seem to balance each other when coupled with the constant small number of spawn produced by the nudibranchs. Whether this case has biological significance is questionable, as it is unclear if spawn would be produced by the emigrating nudi- branchs in the absence of food. Experiment S.J,E.. Both the means-only (Figure 5C) and the normal distribution (Figure 5D) show much the same pattern as S.I,E,: a smooth decline in population from its peak, followed by a more precipitous decline (in means-only case) to a low, constant value (N.,g = 14.8 + 0.1 means-only, 21.2 + 0.5 normal-distribution). Experiment S,J,E.. (Figure 5E, F). This experiment shows much the same pattern as the preceding one except that the population goes to zero in both cases long after the food is exhausted (t, = 24.1 + 0.1 means-only, 32.7 + 0.4 normal-distribution). DISCUSSION The computer model predicts several things about the be- havior of the nudibranch-hydroid system. One prediction is that Cordylophora lacustris will not survive predation by Tenellia adspersa, based on the hydroid/nudibranch growth rates and nudibranch predation rates used in the models. Cordylophora lacustris persists for the longest time under conditions of no adult migration (mmigration or emigration) or larval settlement (experiment S,I,E,) or with only larval settlement (experiment S.J,E,). Once adult immigration is taken into account, either through constant immigration (experiment S.I.E,) or through im- migration only when hydroid remains (experiments S_[,E., and S,J,E..), C. lacustris persists a fraction of the time. This appears consistent with field and laboratory ob- servations. For example, laboratory populations of C. /a- custris colonies initially containing 86.8 + 9.7 polyps persisted for 9.0 + 1.1 days with four juvenile 7. ad- spersa (n = 4). In addition, Turpaeva (1963) demonstrat- ed that a single individual of Tenellia adspersa could de- stroy a colony of the hydroid, Perigonimus megas con- sisting of up to 100 polyps in as short as 24 hours. Another prediction of the simulations is that in all cases nudibranchs will persist for a long time after their food supply is exhausted, even under conditions that most closely mimic field conditions. This has implications for successional change within fouling communities. Not only will the nudibranchs destroy the hydroid colony, but also their presence prevents recolonization by hydroids. C. M. Chester et al., 2000 Maturity Size (mm) micinorD os 1.000 0.100 Survival to Age x (1,) 0 5 10 Page 343 300 7) 250 200 150 100 GGG] WAG GY Don —YVAY A\4\7 nV WA7 W\7 W\7 W7 A\7 AVA WAG yY ); Z 7 Z Z JS ||I[ | W4Uo"g RG Wk MS, gg UMA GG MSS Zs AVAVAIZ ZA # of Eggs - Individual! - Day! 15 20 Da) 30 Age (days) Figure 2 Nudibranch size and fecundity (A) and survivorship data (B) for Tenellia adspersa raised on Cordylophora lacustris at 30%o salinity and 25°C. This may help to explain why hydroids are often ephem- eral and tend to be succeeded by other species. In addi- tion, this suggests that nudibranchs may indirectly affect the successional process by removing hydroids and pre- venting their re-establishment. Tenellia adspersa may also have an effect on this process because of its feeding mechanisms. Tenellia adspersa predominantly feeds by piercing the perisarc and sucking out the tissue contents. The perisarc is not eaten and typically remains behind. Although the three-dimensional structure, and hence ef- fects on current flow over the colony, could remain much the same, a healthy hydroid colony, with its polyps intact, will very likely impede or facilitate settling of larvae quite differently than will its skeleton (Standing, 1976). A third prediction is that in the presence of constant larval settlement (experiments S.I,E,, S.I,E,, S.IJE., and Page 344 8 pte | > 6 Zz eo) 2 Boe an3 ce a2 tt ] 0 Volume of Stolon Eaten (mm?*day') Ny) lied 9 223 ie Ae Nudibranch Size (mm) Figure 3 Predation rates for Tenellia adspersa feeding on Cordylophora lacustris in terms of number of polyps eaten per day (A) and volume of stolon eaten per day (B). The results of a Tukey HSD are presented as an asterisk if significantly different, and lines if not significantly different at a = 0.05 level. S.[E.), nudibranchs will be present in an area no matter what the status of the hydroid colony. The nudibranch growth studies revealed that newly metamorphosed slugs persist for several days before visibly feeding on the hy- droid. If there is no other food supply available, it is unlikely these nudibranchs will survive to adulthood. However, the presence of a steady supply of juvenile nu- dibranchs will prevent recolonization by hydroids. The hydroid, Cordylophora lacustris, lives in fresh or brackish water conditions and can tolerate greater fluc- tuations in its habitat than its marine relatives (Fulton, 1962). In the Great Bay Estuary, C. lacustris is only The Veliger, Vol. 43, No. 4 found in lower salinity riverine systems (15—20%oc or less) and does not occur within the Great Bay (Chester, per- sonal observation). Although not performed under a range of salinities, the hydroid growth study tends to sup- port these observations. Higher growth rate in terms of both number of polyps and volume of stolon occurred at 20%c salinity with slower growth occurring at 25%c. Un- der controlled conditions, using defined media, Fulton (1960) grew colonies of Cordylophora lacustris and found that polyps increased exponentially with a doubling time of 3 days. Stolons grew linearly with a growth rate of 0.1 mm/hr (Fulton, 1963). This translates to a change in volume of 1.5 mm/?/day (using a stolon diameter of 0.2 mm [Fulton, 1961]). The present study yielded values higher than Fulton’s (1961) for the hydroid in 20%c. The growth dynamics of Tenellia adspersa presented in this study are similar to those found by Harris et al. (1980) and Rasmussen (1944). However, the generation time and life cycle were shorter than previously observed. This may be a result of varying laboratory conditions. Tenellia adspersa grew exponentially until about the time of the first spawn mass. Growth rates decreased until about the 24th day when growth rates were negative. As with previous studies, fecundity varied greatly among in- dividuals. Tenellia adspersa produces both pelagic lecithotrophic larvae and capsular metamorphic juveniles, so some of the offspring will remain within the hydroid colony. This was accounted for in the simulations. Coupled with the short generation time observed (2—3 weeks), nudibranchs will build up within a colony very quickly as observed in the simulations. This will affect predation rates and ultimately the persistence of the hydroid colony. The taxonomy of Tenellia is confusing. There appears to be at least two species of Tenellia along the Atlantic coast; Tenellia fuscata (Gould, 1870) and Tenellia ad- spersa (Nordmann, 1845). A third name, Tenellia pallida (Alder & Hancock, 1854) is a synonym of T. adspersa (Roginskaya, 1970). Tenellia fuscata possesses a mus- cular hermaphroditic duct consisting of a set of muscular sphincters located in the anterior portion of the oviduct (Chambers, 1934), lacks an armed penis, and lacks cni- dosacs (Marcus & Marcus, 1960). The penis of Tenellia adspersa possesses a cuticular stylet (Chambers, 1934; Marcus & Marcus, 1960) as well as the presence of small cnidosacs at the tips of the cerata (Roginskaya, 1970; Brown, 1980). All of the available evidence, including radula morphology indicates that the name Tenellia ad- spersa is justified (Gosliner, personal communication). The pattern described in these simulations for this spe- cific nudibranch-hydroid association is consistent with the general model that early colonists do not replace them- selves (Connell, 1975). The mechanism for this pattern is the accumulation of predators that consume the colonists and persist long enough to both inhibit re-establishment of the colonists and to facilitate the development of later #Slugs and Spawn # Slugs and Spawn # Slugs and Spawn . M. Chester et al., 2000 6000 5000 4000 3000 2000 1000 180 1200 160 — SS 1000 1402 ¢ 73 «6S 120 S 8g 800 no} n 100 = = 600 0) Ga o & 60 € = 400 Or eS 200 2 2 0 0 On LOm2 030814050 260870) 780 OPO 420° 30° 40° -50' 60) 70, 80 Time (days) Time (days) Amount of hydroid (%) 0 10 20 30 40 50 60 70 80 0 10 20 30 40 50 60 70 80 time, (days) Time (days) 0 iS —— Adults 160 eS a a. Spawn 140 © 5 Hydroid mo} WAG = nN 100 = 3 80 os Sy OS AQ Gk 20 < 0 0 10 20 30 40 50 60 70 80 0 10 20 30 40 50 60 70 80 Time (days) Time (days) Figure 4 Results of discrete-event simulations: (A) S,I,E, means-only, no larval settlement, no immigration, and no emigra- tion, (B) S,I,E, normal distribution, no larval settlement, no immigration, and no emigration, (C) S,I,E, means- only, continuous settlement, no immigration, and no emigration, (D) S,I,E, normal distribution, continuous settle- ment, no immigration, and no emigration, (E) S,l,E, means-only, continuous settlement, no immigration, and emi- gration during starvation, (F) S,,E, normal distribution, continuous settlement, no immigration, and emigration during starvation. Page 345 35 30 S 250 e 20 2 as 15 & Ons = 5 0) Amount of Hydroid (%) Amount of Hydroid (%) The Veliger, Vol. 43, No. 4 # Slugs and Spawn O° 10% 2030840550 260 70ms0 Time (days) Smee AGUIES a aan pawn Hydroid # Slugs and Spawn O10) 20° 30% 40550) 607 705480 Time (days) # Slugs and Spawn 0 10) 205 30) 40° 5057605707 180 Time (days) mo) Tc 5 5 mo} Tc cal > x a5 oe oH cs) S iS e 3 = © S E = << < 0 10) 20° 30° 40> S50 COM 7Oms6 Time (days) 120 5) x= 2 100 4 x ee z 5 a. 80 iS) a8 5 col witch (6) ee Ge oH ° an 2 ee a S 4 2 ie & 3 = < ED < 0 0 0 10 20 30 40 50 60 70 80 Time (days) ——"Adults | > a —— — Spawn So oS : SS =o Hydroid [> 4 © aS) as) 5 S 3 Sa. as = = = £ 8 28 iS = S =) WY 5 E * ie < < 0 0 10 20 30 40 50 60 70 80 Time (days) Figure 5 Results of discrete-event simulation: (A) S,I,E, means-only, settling only when food is present, no immigration, and emigration when starving, (B) S,I,E, normal distribution, settling only when food is present, no immigration, and emigration when starving, (C) S,I,E,, means-only, continuous settlement, immigration, and emigration, (D) SIE, normal distribution, continuous settlement, immigration, and emigration, (E) S,1,E,, means-only, continuous settlement, immigration when food is present, emigration increased during starvation (F) S,,E,, normal distribution, continuous settlement, immigration when food is present, emigration increased during starvation. C. M. Chester et al., 2000 Page 347 Table 2 Results of the simulation experiments. S = larval settlement, I = immigration, E = emigration, f = fed, s = starved, y = yes, n = no, | = 1%/day, 10 = 10%/day. All data shown are the average of 50 runs. Means-only runs were started with enough food for the 15 nudibranchs for 1 day, normal-distribution runs with enough for seven nudibranchs for | day. All t, values are significant (P < 0.0001). 14.8 (0.1) Normal-distribution case S I E Means-only case Exp f Ss f s f s ty = 0 IN Nena SJE, n n on on on _n 49.2 (0.4) 4643.2 (55.1) SREY. Dn on nn 4616.1 (60.1) 34.0 (0.0) SHEESEyny, 0) mn) 10 3221.2 (42.4) 14.9 (0.2) SIE, y n on on on 10 48.4 (0.2) 3240.0 (35.0) SHEEREyEY oy oy Ll 1 881.5 (8.8) 735.3 (7.4) SHE Sy SY ol) 228.6 (7.3) SPeeyeene yo om 1 10) 2707) 24051 7) successional stage species (Standing, 1976; Harris et al., 1985). Predation on grazers within early colonists could lead to persistence of early colonists to inhibit succession or at least alter the successional sequence to other species (Standing, 1976). Acknowledgments. We wish to thank L. Dyer, S. Iskra, S. Mul- liken, M. Nyberg, B. Piel, L. Rodriguez, EF Rotman, K. Sardi, K. Siedl, and C. Williams for help with the hydroid and nudibranch cultures, and J. Beal for help with the computer simulation. Drs. M. Litvaitis, J. Haney, A. Kuzirian, E. Tillinghast, and two anon- ymous reviewers provided invaluable comments on earlier drafts. Portions of the lab studies were supported by a Grant-in-Aid of Research from Sigma-Xi to CMC. LITERATURE CITED ANDERSON, S. D., D. L. WESTBROOK, M. SCHMILL, A. CARLSON, D. M. Hart & P. R. COHEN. 1994. Common Lisp Analytical Package Users Manual. University of Massachusetts: Am- herst, Massachusetts. BANKS, J., J. S. Carson, HI & B. L. NELSON. 1996. Discrete- Event Simulation. 2nd ed. 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ZAR, J. H. 1996. Biostatistical Analysis. Prentice Hall: New York. 120 pp. The Veliger 43(4):349-—353 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 Predation on the Apple Snail, Pomacea canaliculata (Ampullariidae), by the Norway Rat, Rattus norvegicus, in the Field YOICHI YUSA, NAOYUKI SUGIURA Anp KATSUYA ICHINOSE Kyushu National Agricultural Experiment Station, Nishigoshi, Kumamoto, 861-1192 Japan; e-mail: yusa@knaes.affrc.go.jp Abstract. Aspects of predation on the apple snail, Pomacea canaliculata, an important pest of rice, by the Norway rat, Rattus norvegicus, were studied near a drainage canal in southern Japan. Most of the shells attacked by the rats were broken in a characteristic way, from the outer edge of the last whorl up to the attachment site of the columellar muscle. The rats ate all the flesh of the snails except for the albumen gland of the female. They preferred Pomacea canaliculata to another freshwater snail Semisulcospira libertina. They preferred large Pomacea snails to small ones but showed no preference between the sexes of the snails. The numbers of snails attacked within 12 hours varied from O— 12 per rat hole, and predation was almost always nocturnal. INTRODUCTION The apple snail, Pomacea canaliculata (Lamarck, 1804), was introduced into Asia from South America mainly in the 1980’s as food for humans (Halwart, 1994a, b; Wada, 1997). However, its commercial value was soon lost, and abandoned snails became wild in many Asian countries including Cambodia, China, Indonesia, Korea, Japan, Laos, Malaysia, Taiwan, Thailand, Papua New Guinea, the Philippines, and Vietnam (Halwart, 1994a, b; So- mony, 1998). Areas infested by this snail in these coun- tries are still expanding (Wada, 1997), and other neigh- boring countries (e.g., India and Australia) are threatened with invasion (Baker, 1998). P. canaliculata feeds vora- ciously on rice seedlings and has now become one of the most important pests of rice in Asia. Due to difficulties in applying chemical and cultural techniques to control this snail, natural enemies have been suggested as biological control agents (Kondo & Tanaka, 1989; Halwart, 1994a, b; Wada, 1997). In addition, proper understanding of the population dynamics of the snail in the field depends in part on identifying the influence of predation on snail numbers. Until now, several wild ani- mals have been observed to feed on Pomacea canalicu- lata (Hamada & Matsumoto, 1985; Kondo & Tanaka, 1989; Ozawa et al., 1989; Halwart, 1994a, b; Chanyapate, 1997; Suzuki et al., 1999), but few of these predators attack adult snails (but see Ozawa et al., 1989). No at- tempts have been made to quantify predation on this snail by wild animals in natural habitats. Rodents have been observed feeding on P. canalicu- lata in Thailand (Chanyapate, 1997) and the Philippines (Almazan et al., unpublished data). In March 1998, piles of broken shells of this snail, apparently attacked by a species of rodent, were found near a drainage canal in southern Japan. Using video recordings, we identified the rodent to be the Norway rat, Rattus norvegicus (Berken- hout, 1769). In this paper, we report aspects of this rat’s predatory behavior and discuss the influence of its pre- dation on snail numbers. MATERIALS AnD METHODS Study Area The canal along which we found many broken shells of P. canaliculata was in Shichijo, Kumamoto Prefecture, Kyushu Island, southern Japan (130°50’E, 33°10’N). The water in the canal was 1.5 m wide and approx. 0.3 m deep. Both sides of the canal were lined with concrete walls (0.5 m high), above which soil banks extended 2 m farther. The bottom of the canal was also lined with concrete, but sand and pebbles completely covered this. Four rat holes, two on each side of the canal, were dug into the soil banks just above the concrete walls of the canal. The two holes on each side were approx. 1.5 m apart from each other and 5—7 cm in diameter. Near the opening of each hole, broken shells and opercula of P. canaliculata were discarded on the ground, as well as those of other snail species. Collection of Snails Apparently newly discarded shells and opercula of ap- ple snails and other snail species were removed from the ground near the rat holes on 10 March 1998. After that, the rat holes were observed twice per day, once in the morning (between 0620 and 0730) and once in the even- ing (between 1800 and 1900) for 10 days to examine the frequency of predation per rat hole. When newly discard- ed shells or opercula were found, they were brought to the laboratory for species, size, and sex determination. Page 350 On 21 March, after the collection of discarded shells near the rat holes, we collected living snails up- and downstream in the canal near the holes (15 m along the stream and 1.5 m across). First, we collected all the snails on the surfaces of the canal (both on the concrete walls and on the sediment) within this area. We then collected snails buried in the sediment by sieving this material through hand nets with 5 mm mesh. Shell Measurements Shell heights were measured for all the Pomacea shells collected, both near the rat holes and in the canal. The length and width of broken shell parts were also measured for the individuals collected near the rat holes. Length was measured as the longest curvilinear distance of the broken part of the shell along the whorl. Width was mea- sured vertical to the direction of the length (hence parallel to the shell axis), and measured three times along the whorl (twice near both ends and once in the middle) for each shell and then averaged. Sexual dimorphism occurs in this apple snail: mature females have similar opercula and shell shapes to those of juveniles, but the opercula of mature males have re- flected edges, and their shells have extended outer edges (Cazzaniga, 1990; Estebenet, 1998). Thus, sexes can be identified on the morphology of the shell or operculum after snails reach sexual maturity. Amongst a subsample of snails collected in the canal, all males 20 mm or larger in shell height had reflected operculum edges and fully developed testes (12 individuals dissected), and most (89%; 16/18) of the females in this size had well-devel- oped albumen glands (cf. Kaneshima et al., 1986). Hence, for the snails collected from the canal, those with shell heights of 20 mm or larger were sexed, based on the morphology of the shell and operculum. Sexes of the attacked snails were identified using only their opercula, since the outer edges of their shells were lost in most cases. Shell height for these snails also had to be estimated from the size of the operculum. The re- gression of operculum length on shell height was highly significant among snails in the canal (shell height = 1.458 x operculum longitudinal length, 7? = 0.968, n = 88, P < 0.0001). Using this regression equation, an operculum length of 13.7 mm corresponded to a shell height of 20 mm. Sex was thus determined for snails with discarded opercula of this size or larger. RESULTS Characteristics of Attacked Snails We collected 196 discarded shells of P. canaliculata near the four rat holes. Most of the discarded shells were damaged and empty, whereas 19 shells (10%) were intact. Damaged snails were significantly larger (mean + SD shell height = 21.5 + 5.4 mm, n = 177) than intact ones The Veliger, Vol. 43, No. 4 Figure | Shell of Pomacea canaliculata broken by the Norway rat, Rattus norvegicus. Shell height is 29 mm. (11.9 + 5.6 mm; t = 7.35, P < 0.0001). Most of these latter snails were still alive when collected. The rats damaged the shells of apple snails in a char- acteristic way (Figure 1). All the damaged shells were broken from the outer edge along the whorl, and usually far less than one whorl was broken. The shapes of the broken parts were similar, irrespective of the shell height. Thus, there were significant correlations between shell height and the curvilinear length of the broken part along the whorl (n = 175: two cases were excluded in which the shell was broken so heavily that measurements were impossible, r = 0.726, P < 0.0001; Figure 2a), and be- tween shell height and the width of the broken part (r = 0.877, P < 0.0001; Figure 2b). The flesh of the snails was completely eaten in most cases, except for the al- bumen gland of the female, which was discarded near the rat holes along with the broken shells. Differences between Attacked Snails and Snails in the Canal Shells of three species of freshwater snails were found near the rat holes (Table 1). The proportion of P. canal- iculata was highest (88.7%), followed by the pleurocerid Semisulcospira libertina (Gould) (10.9%). Only one spec- imen of the vivipariid Cipangopaludina chinensis laeta (Martens) was found near the rat holes (0.5%). On the other hand, among snails living in the canal, the propor- tions of P. canaliculata (54.4%) and of S. libertina (45.5%) were similar. The proportion of these two major species varied significantly between the two sites (x? = Y. Yusa et al., 2000 £ on =) (=) N S Length of broken part (mm) 0) 10 20 30 40 Page 351 Width of broken part (mm) 0 10 20 30 40 Shell height (mm) Figure 2 Relationships of shell height to the length (a) and width (b) of the broken parts of shells of Pomacea canaliculata. 91.86, P < 0.0001) including all the snails collected. Al- though only snails that seemed newly discarded were col- lected at the start of the observations, some of these snails might have been there for a long time. If so, this differ- ence between sites might have been due to changes in species composition over time rather than preference of rats. This possibility, however, could be refuted because the proportion of species varied significantly between snails in the canal and those near rat holes collected over the 10-day observation period (x7 = 18.89, P < 0.0001). The discarded Pomacea snails (mean + SD shell height = 20.5 + 6.1 mm, n = 196) were much larger than living conspecifics in the canal (10.6 + 5.1 mm, n = 612; t = 20.23 after log-transformation, P < 0.0001; Figure 3). Among the snails living in the canal, those on the sur- faces (concrete walls and sediment) were larger (12.4 + 5.8 mm, n = 56) than those buried in the sediment (10.4 + 5.0 mm, n = 556; t = 2.85 after log-transformation, P < 0.01). The shells discarded near the rat holes were still much larger than the snails on the canal’s surfaces (t = 10.03 after log-transformation, P < 0.0001). This last dif- Table 1 Numbers of snails attacked by the Norway rat and living in the nearby canal. Snail species Pomacea_ Semisulcospira Cipangopaludina Site canaliculata libertina chinensis Rat holes (total) 196 24 1 Rat holes (collected over 10 days) 54 12 0) Canal 612 511 1 ference between sites was still significant even if snails collected at the start were excluded from the analysis (t = 5.49 after log-transformation, P < 0.0001). The number of opercula found near the rat holes (n = 175) was similar to that of the empty shells without oper- cula (n = 184). The sex ratio estimated from the opercula should thus reflect the sex ratio of the attacked snails. The numbers of male and female opercula near the rat holes were 71 and 63, respectively, where operculum length was 13.7 mm or longer (corresponding to a shell Rat holes @ 10 days C) at start 200 surfaces O buried 150 No. of snails 100 0 10 20 30 40 Shell height (mm) Figure 3 Size distributions of Pomacea canaliculata discarded near rat holes (upper) and those living in the canal (lower). Note that scales on the ordinate axis are different in the two graphs. The Veliger, Vol. 43, No. 4 No. of snails attacked 9 20 Mar: ti) 12. 13i WAS aeelGnen le, aus Date Figure 4 Numbers of Pomacea canaliculata attacked by rats, recorded at 12-hour intervals. Different symbols indicate different rat holes. Dark areas indicate night-time. height of 20 mm or higher, including two intact males). The sex ratio for the discarded snails was not significantly different from the sex ratio of the snails collected in the canal (28:21, x? = 0.25, P = 0.62). This difference was also nonsignificant when the opercula collected at the start were excluded from the analysis (xy? = 0.32, P = 0.57). Frequency of Predation The number of newly discarded shells recorded at 12- hour intervals varied between rat holes, sampling times, and days, from O (in most cases) to 12 (Figure 4). The average number of discarded shells per day per hole was 1.4. Attacks were much more frequent at night (n = 52) than in the day (n = 2; P < 0.0001, binomial test). DISCUSSION Most of the attacked shells of P. canaliculata were bro- ken from the outer edge, and the size of the broken part was proportional to shell height, usually far less than one whorl. This pattern of damage suggests that the rats peeled off the shells with their teeth, until they reached the attachment site of the columellar muscle. A laboratory observation on the behavior of the rat confirmed this point. In addition, the rat avoided the albumen gland of the female snail. This was probably due to its bitter taste (based on the authors’ personal tasting). In Colombia, the snail kite, Rostrhamus sociabilis sociabilis, is also known to discard the albumen gland of Pomacea chemnitzi when feeding (Snyder & Kale, 1983). The rats attacked proportionally more P. canaliculata than S. libertina in the canal. The rats also preferred larg- er P. canaliculata to smaller ones. Feeding costs versus benefits may be relevant to these preferences. P. canali- culata is generally larger in size and has a thinner shell than S. libertina. Rats would therefore gain more energy with far less effort by preying upon P. canaliculata. The broken area of the Pomacea shell is also a square function of shell height (since both length and width of the de- stroyed part are correlated with shell height), whereas body weight increases as a cubic function of length (at least in Pomacea dolioides: Bourne & Berlin, 1982). Thus, larger snails are likely to provide more benefit per effort for a rat. There was relatively little difference in size between male and female P. canaliculata in this study, and the rats showed no preference. Many wild animals have been reported as predators of P. canaliculata, including leeches, aquatic insects, crabs, fishes, birds and mammals (Hamada & Matsumoto, 1985; Kondo & Tanaka, 1989; Ozawa et al., 1989; Halwart, 1994a, b; Chanyapate, 1997; Suzuki et al., 1999). In most cases, however, the impact of their predation is unknown, and few predators are known to feed on adult snails (with the exception of leeches; Ozawa et al., 1989). This study has shown that Norway rats prefer large P. canaliculata, and has suggested that at least on some occasions, pre- dation rate may be quite high (e.g., 12 shells found at one rat hole in one night). In addition, laboratory observations showed that individual Norway rats can consume 68—72 adult snails (shell height = 20-30 mm) per day (n = 3; Yusa, personal observation). Considering its potentially high predation rate, preference for large snails, and wide distribution, the Norway rat seems likely to be one of the most important predators of the apple snail in the field. In fact, predation by rats (species not identified) has been noted as the main cause of the mortality of the apple snails in paddy fields at the International Rice Research Institute, the Philippines (Almazan et al., unpublished data). However, strong recommendation of this rat as a biological control agent is very unlikely, given that the rats themselves are serious pests of rice, and they can also be hosts of various bacteria or invertebrates parasitic to humans, including the rat lung worm, Angiostrongylus cantonensis, which uses P. canaliculata as an interme- diate host and causes eosinophilic meningoencephalitis (Nishimura & Sato, 1986). Acknowledgments. We wish to thank Dr. S. Urano for help in identifying the rat species, Ms. J. Murakami for help in measur- ing snails, and Drs. T. Wada and G. H. Baker for discussion and critical reading of the manuscript. LITERATURE CITED BAKER, G. H. 1998. The golden apple snail, Pomacea canali- culata (Lamarck) (Mollusca: Ampullariidae), a potential in- vader of fresh water habitats in Australia. Pp. 21—26 in Sixth Australian Applied Entomological Research Conference, Volume 2. The University of Queensland. Bourne, G. R. & J. A. BERLIN. 1982. Predicting Pomacea do- lioides (Reeve) (Prosobranchia: Ampullariidae) weights from linear measurements of their shells. The Veliger 24: 367-370. CaZZANIGA, N. J. 1990. Sexual dimorphism in Pomacea canali- Y. Yusa et al., 2000 culata (Gastropoda: Ampullariidae). The Veliger 33:384— 388. CHANYAPATE, C. 1997. The golden apple snail, Pomacea canal- iculata research and management in Thailand. 4 pp. in Pro- ceedings of the International Workshop on Ecology and Management of the Golden Apple Snail in Rice Production in Asia. The International Rice Research Institute, Swiss Agency for Development and Cooperation & Department of Agricultural Extension, Thailand. ESsTEBENET, A. L. 1998. Allometric growth and insight on sexual dimorphism in Pomacea canaliculata (Gastropoda: Ampul- lariidae). Malacologia 39:207—213. Hacwart, M. 1994a. The golden apple snail Pomacea canali- culata in Asian rice farming systems: present impact and future threat. International Journal of Pest Management 40: 199-206. Hacwart, M. 1994b. Fish as Biological Agents in Rice. Margraf Verlag: Weikersheim, Germany. ili + 169 pp. Hamaba, Y. & T. Matsumoto. 1985. Apple snails in Kumamoto Prefecture. Kyushu-no-kai 24:5—12 [in Japanese]. KANESHIMA, M., S. YAMAUCHI & K. HIGA. 1986. Sexual maturity of the apple snail, Ampullarius insularus. Proceedings of the Association for Plant Protection of Kyushu 32:101—103 [in Japanese]. Konpbo, A. & E TANAKA. 1989. An experimental study of pre- dation by the larvae of the firefly, Luciola lateralis Mot- schulsky (Coleoptera: Lampyridae) on the apple snail, Po- Page 353 macea_ canaliculata Lamarck (Mesogastropoda: Pilidae). Japanese Journal of Applied Entomology and Zoology 33: 211-216 [in Japanese with English abstract]. NISHIMURA, K. & Y. SATo. 1986. Natural infection with Angios- trongylus cantonensis in Ampullarius canaliculatus (La- marck) in the Ryukyu Islands, Japan. Japanese Journal of Parasitology 35:469—470. Ozawa, A., T. MAKINO & S. IsiGAMI. 1989. A predator of adults of the apple snail, Pomacea canaliculata (Lamarck), a spe- cies of leeches, Whitmania pigra. Proceedings of the Kanto- Tosan Plant Protection Society 36:214 [in Japanese]. SNYDER, N. EF R. & H. W. Ka te, II. 1983. Mollusk predation by snail kites in Colombia. The Auk 100:93-97. Somony, C. 1998. A short review of the golden apple snail in Cambodia. 4 pp. in Proceedings of the International Work- shop on the Integrated Management of the Golden Apple Snail in Rice Production in Vietnam, FAO & Plant Protec- tion Department, Vietnam. SuzuKI, Y., K. MryAmoto, M. MatTsumuraé, K. ARIMURA & F TUBIANO. 1999. Enemies of the golden apple snail Pomacea canaliculata juveniles in paddy fields. Kyushu Agricultural Research 61:83. [in Japanese] Wapa, T. 1997. Introduction of the apple snail Pomacea canal- iculata and its impact on rice agriculture. Pp. 170-180 in Proceedings of International Workshop on Biological Inva- sions of Ecosystems by Pests and Beneficial Organisms. Na- tional Institute of Agro-Environmental Sciences, The Min- istry of Agriculture, Forestry and Fisheries, Japan. The Veliger 43(4):354—366 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 Three New Species of Dorid Nudibranchs from Southern California, USA, and the Baja California Peninsula, Mexico SANDRA V. MILLEN Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver, B.C., Canada, V6T 1Z4 AND HANS BERTSCH Research Associate, California Academy of Sciences, Golden Gate Park, San Francisco, California, 94118, USA! Abstract. Three new species of dorid nudibranchs, one phanerobranch Trapania goslineri, and two cryptobranchs, Peltodoris mullineri and Peltodoris lancei, have been found in the Gulf of California and nearby Pacific waters of the Baja California peninsula, Mexico, and southern California, USA. Trapania goslineri can be distinguished by its white body with large black blotches and yellow tips. Peltodoris mullineri is a large, yellow to golden orange dorid, with speckled brown blotches. Peltodoris lancei is a large, dark orange dorid, which has small white-tipped papillae and a very depressed body. INTRODUCTION The opisthobranch fauna of Baja California, on the Pa- cific side and in the Gulf of California, is less known than the fauna of the more temperate zone farther to the north. Some indication of the rich diversity can be found in the opisthobranch chapter of Keen (1971), its update by Skoglund (1991), and in Bertsch (1989). Studies of the Pacific coast of the peninsula have noted the presence of several undescribed species (Gosliner et al., 1985; Bertsch, 1991b). In the waters of the Gulf of California, an ongoing survey of the ecology and abundance of opis- thobranch mollusks was initiated by Hans Bertsch in the early 1980s. This study has been primarily centered at Bahia de los Angeles, Baja California, Mexico. During the course of this study, a number of rare and undescribed nudibranchs have been discovered (Bertsch, 1991a, 1995a, b). Some of these species have since been de- scribed (Gosliner & Behrens, 1986; Gosliner et al., 1999). This paper describes three additional species: Peltodoris lancei from the Gulf of California, Trapania goslineri, which is found on both coasts of the Baja California pen- insula, and Peltodoris mullineri from the outer coast of southern California and Baja California. ' National University, 192 Imperial Beach Boulevard, #A, Im- perial Beach, California 91932, USA. SYSTEMATICS Family GONIODORIDIDAE H. & A. Adams, 1854 Genus Trapania Pruvot-Fol, 1931 Trapania goslineri Millen & Bertsch, sp. nov. (Figures 1A, 2A—E 3) Trapania sp. Behrens, 1983: 19; Bertsch & Kerstitch, 1984: 264; Bertsch, 1989:63; Bertsch, 1996. Trapania sp. 1 Behrens, 1991:47 Etymology: This species is named for our friend, the pro- lific opisthobranch researcher and photographer, Terrence Gosliner of the California Academy of Sciences. Material examined: Holotype: CASIZ 118571, Punta la Gringa (29°02.56’N, 113°32.29'W), Bahia de los Angeles, Baja California, Mexico, 29 June, 1987, collected by T. Gosliner. Paratypes: CASIZ 112214, 2 specimens, one dissected, from the type locality, 29 June 1987, collected by H. Bertsch. Other material: Not deposited, 1 specimen, dissected, Cuevitas, Bahia de los Angeles, 26 May 1995, collected by H. Bertsch. Not deposited, 1 specimen and spawn mass, Bahia Tortugas, 6 October, 1996, H. Bertsch. Not deposited, 1 specimen, dissected, Punta Don Juan, Bahia de los Angeles, 5 October, 1984, H. Bertsch. External morphology: The soft body has a limaciform shape with the mantle reduced to two pairs of posteriorly S. V. Millen & H. Bertsch, 2000 curved lateral processes, one pair beside the gills and one lateral to each rhinophore, as is typical for the genus (Fig- ure 1A). The measurements of a typical preserved spec- imen are 8.5 X 2.5 X 3 mm (1 X w X h). The largest living specimen had a length of 15 mm. The non-retrac- tile rhinophores have no sheaths and 9-10 leaves. There are three non-retractile, bipinnate gills, one medial and two lateral, all anterior to the anus. The anus is on a small papilla, and the renal opening is above it, between the right and central gills. The head has digitiform oral ten- tacles extending antero-laterally. The narrow foot is ex- panded anteriorly into short, recurved, propodial tentacles and is bilabiate on its anterior edge. The ground color is white with large, black oval spots and streaks (Figures 1A, 2A, B). There is a V-shaped, wide black mark on the head from in front of the rhin- ophores extending onto the base of the oral tentacles. There is a wide, longitudinal stripe extending from behind the rhinophores half way back to the gills, which may be confluent with spots in front of the gills. The branchial and rhinophoral processes have black streaks originating on the body and extending out their dorsal sides. Their tips are golden orange. The oral tentacles are white or spotted basally, golden orange distally. The rhinophores and propodial tentacles are white basally with golden or- ange distally. The three bipinnate gills are translucent white, sometimes with large black spots near their bases, with golden orange tips. Large oval to round black spots are irregularly distributed on the sides and midline. The tail is golden orange with a medial streak of the same color. Ventrally, the head and foot are white. There are minor variations from this pattern. One specimen had white instead of golden orange on the tail, branchial and rhinophoral processes, and oral tentacles, but not the gills and rhinophores. Several specimens had smaller, more numerous black spots. Another specimen had golden or- ange extending most of the length of the lateral processes. Internal anatomy: There are simple labial glands ventral to the mouth opening. The very short, ringlike buccal tube leads to a large, round, sessile buccal bulb with no projecting radular sac. The suctorian crop has transverse muscles but no median longitudinal muscle division. In- side there are two triangular jaw plates with a triangular patch of pointed rodlets up to 27 wm in length (Figure 2C, D). The radular formula is 37—41 (1.0.1). The teeth are elongate plates with a recurved distal edge bearing denticles. There are two small outer denticles, the second of which is sometimes subdivided, at the base of the long narrow cusp which is two-thirds of the distance to the outside of the tooth. The inner edge has seven to nine intermediate denticles varying in size, the longest of which is half the length of the cusp and just outside of center. In addition, there are a number of minor inter- mediate serrations which are only visible using scanning electron microscopy (SEM) (Figure 2E, F). Posteriorly, Page 355 on either side of the esophagus are two small, round, salivary glands. The esophagus is long and tubular. The striated stomach is buried in the confluent digestive glands and possesses a large, round posterior caecum, which is muscular and free of the digestive glands. The narrow, tubular intestine leaves the stomach at its junction with the caecum and travels anteriorly to the anterior edge of the digestive gland, then curves abruptly to the right and posteriorly to end on a small anal papilla. The central nervous system has fused cerebro-pleural ganglia with large, sessile eyes. The ventral pedal ganglia are large and round. No visceral ganglia were seen. On the ventral side of the pharynx, the two large, round buc- cal ganglia are close together. The triaulic reproductive system is illustrated in Figure 3, excluding the ovotestis, which lies in lateral rows along the top and sides of the digestive glands. The round fe- male acini lie on either side of the soft, tubular male acini. Ductules join them to the median longitudinal hermaph- roditic duct. This duct is free of the digestive glands for a short distance on the anterior right, then it loops dor- sally alongside the hermaphroditic ampulla and enters it one-third of the way from its distal end. The large, oval ampulla lies on the female gland mass. At its distal end the ampulla bifurcates into a vas deferens and a short oviduct, which unites with the insemination duct and en- ters the female gland mass. The vas deferens swells into a long, tubular prostatic portion, which bends once and constricts distally before a short muscular section, ter- minating in a small muscular penis at the common atrium. The penis is armed with several rows of slender, slightly curved spines. The vagina is tubular, wider at the com- mon atrium. The narrow portion bends in an S shape and extends dorsally, then gives off the insemination duct 0.1 mm before the bursa copulatrix. The bursa copulatrix is large, spherical, and dark with sperm. The narrow, tubular insemination duct runs alongside the smaller, oval recep- taculum seminis, which is inserted on it by a short duct. The insemination duct continues to join with the oviduct and forms a fertilization chamber before entering the fe- male gland mass. The female gland mass is round and compact. It has a small granular albumen gland, a highly folded mucus gland, and a distal, saccate membrane gland. It exits ven- tral to the vagina and penis in the common atrium. The genital atrium is located far anterior, on the animal’s right side, below and slightly posterior to the base of the rhin- ophore process. ° Natural history: This species is found in the intertidal and shallow subtidal and is very rare. An ongoing, 6-year study by Hans Bertsch in Bahia de los Angeles shows that this species makes up less than 0.2% of the opistho- branch population. Fourteen specimens were observed, the majority on an orange sponge. This genus is thought to feed on kamptozoans which are often attached to Page 356 The Veliger, Vol. 43, No. S. V. Millen & H. Bertsch, 2000 sponge surfaces (Picton & Morrow, 1994). Individuals ranged in length from 8-15 mm (average 10.3 mm) and were found in January (1), March (2), May (3), June (1), July (1), September (4), October (1), and December (1). Egg masses were found twice, in March and October. The spawn was a narrow, white, upright ribbon, of 1% coils, 5-6 mm in diameter, containing approximately 2000 eggs. There was one egg per capsule (Figure 2B). These data are insufficient to clearly describe the life cycle of this species; however, it appears to be more common in spring and summer than in the colder months. This spe- cies has been found off the Baja California peninsula on the Pacific side from Isla Cedros (Behrens, 1983) to Cabo San Lucas (Bertsch & Kerstitch, 1984) and in the Gulf of California at Bahia de Los Angeles and Sonora (Bertsch, 1991b). Discussion: There are at present 20 described species of Trapania worldwide, two additional undescribed species listed by Gosliner (1987), and one undescribed species from the Pacific side of Costa Rica. Sixteen of the de- scribed species have been discussed by Rudman (1987) with the exception of 7. dalva Marcus, 1972, and three recently described species: T. luquei Ortea, 1989, and T. orteai Garcia-Gomez & Cervera, 1989, and T. hispalensis Cervera & Garcia-Gomez, 1989, both in Cervera & Garcia-Gomez (1989). Most of these species can be easily separated by color from Trapania goslineri sp. nov. The four species which share a white ground color with black markings and yellow tips or streaks are compared in greater detail in Table 1. These species all have pointed jaw rodlets, a semi-serial receptaculum seminis, and sim- ple hooks on the penis. Trapania japonica (Baba, 1935) differs from T. goslineri externally by its much smaller, browner spots which are absent from the lateral process- es. Internally it has unique radular teeth, which have an almost central cusp with 9-12 outer and 12—24 inner den- ticles. Trapania luquei can be distinguished externally from T. goslineri by the yellow spots on the body, yellow patch on the head, and reticulating rather than solid pat- tern of black blotches. Internally it has a much larger cusp on the teeth with only one outer denticle and up to 14 inner denticles. The reproductive system is not known. Trapania toddi Rudman, 1987, also has a reticulating pat- tern to its black blotches. Internally it has similar teeth to T. luquei. Trapania velox (Cockerell, 1901) can be easily distinguished by its color pattern of five black lines in- Page 357 stead of oval spots, and it has a longer tail. Internally T. velox has fewer rows of teeth (22—32 versus 37—41) with one or no denticles external to the cusp instead of two. The bursa copulatrix has a serial arrangement rather than the short common duct found in T. goslineri. Trapania velox is the only species whose range is partly sympatric to that of T. goslineri. Its range is primarily north of T. goslineri, from Hazard Canyon, San Luis Obispo County, California (Roller & Long, 1969). There is a slight dis- tributional overlap in the southern portion of its range, in the region of Isla Cedros and Bahia Tortugas, Baja Cal- ifornia (Behrens, 1991). Family DIsCODORIDIDAE Bergh, 1891 Genus Peltodoris Bergh, 1880 Peltodoris mullineri Millen & Bertsch, sp. nov. (Figures 1B, 4A—F 5) Peltodoris sp. Behrens, 1980: 102—103. Peltodoris sp. Lance, 1983:87; Behrens & Henderson, 1981: 128; Behrens, 1996. Peltodoris sp. 1. Behrens 1991:69; Behrens 1997:13. Etymology: This species is named for the opisthobranch researcher and extraordinary photographer, Dave Mulli- ner of San Diego, California. Material examined: Holotype: CASIZ 074262, Isthmus Cove, Catalina Island, California, 19 October 1985. Coll. T. Gosliner. Paratype: CASIZ 74651, 1 specimen, dis- sected, Diablo Cove, Santa Cruz Island, California, 10— 17 m, 23 August, 1984, collected by R. Van Syoc. Other material: CASIZ 069302, 1 specimen, dissected, Fisher- man’s Cove, Catalina Island, 14 December, 1979, col- lected by David Behrens. Not deposited, 1 specimen, dis- sected, San Clemente Island, 2 November 1985, collected by R. McPeak. External morphology: This large dorid species reaches up to 70 mm in live length and has a depressed oval shape and rigid texture (Figure 1B, 4A). A typical preserved specimen measured 40 < 29 X 10 mm (1 X w X h). The large mantle overhangs the foot by 8—10 mm all around, and the foot is 12 mm wide. The sides are very short. The mantle is covered with small, even, densely set, rounded tubercles which are supported by spicules which protrude from the corners (Figure 4B). The rhinophores Figure | Photographs of living animals. A. Trapania goslineri, Millen & Bertsch, sp. nov. Photo by Jeff Hamann. Specimen from San Ignacio Bay, January, 1998; not collected. B. Peltodoris mullineri, Millen & Bertsch, sp. nov. Photo by Jeff Hamann. Specimen from Point Loma, July 1979; not collected. C. Peltodoris lancei, Millen, sp. nov. Photo by Michael D. Miller, Holotype from Bahia de los Angeles, June 1996 (CASIZ 110807). Page 358 The Veliger, Vol. 43, No. 4 Figure 2 A. Trapania goslineri, Millen & Bertsch, sp. nov. with slightly smaller spots. Specimen from Punta la Gringa, March 1996. B. Trapania goslineri, Millen & Bertsch, sp. nov. with spawn. Specimen from Bahia Tortugas, October 1996. C. Jaw. Scale bar = 100 xm. Specimen from Bahia de los Angeles, May, 1995. D. Jaw denticles. Scale bar = 20 pm. Specimen from Bahia de los Angeles, May, 1995. E. Radular teeth. Scale bar = 10 pm. Specimen from Bahia de los Angeles, May 1995. FE Radular teeth. Scale bar = 10 pm. Specimen from Bahia de los Angeles, October, 1984. S. V. Millen & H. Bertsch, 2000 Page 359 | is s &) 5 3 = ‘3 53 | 2 5 5 mele c ut ay io] Se ee Sida tLe 1 a Z a < & o z 7 & | Se rc beat + re es Celg Baier ie g 2 SS oak 828 gE Bee LN Bel ee oe for a Say ein x a coo) sale @ 3 2\y sa) a) Q ee ee as fog) N Ss nN o z Oe : > | 2 © is) ; e/3s 9 oon ie) O28 Oo oO ° 1mm BD 2 Stan 2 Ag 50 = = |) een fe, 3 e : 3 e|g2s8 ee Ss 26 Figure 3 = Oz — 3 = 5 E 5 Reproductiv =I e system of Tra . 2 ais) ; pania gosli i Mi 2S 0 s ra sp. nov. Scale bar = 1 mm. Speci goslineri, Millen & Bertsch 3 ef 1a a ae Te fen eS May. 1 - Specimen from Bahia d 2 ) a me oO 2 SL y, 1995. Key: bc = b ia de los Angeles = Soe ose Be OB ee fe) Es ursa copulatrix, fgm = Semele 5) Pr |i ees aoe 8S oF a mass, ha = hermaphroditic , fgm = female gland > 4) Bo 2 Sga.3 LON 2 Ee od ye hd = hermaphroditic duct zg o/s? gs fio 278 8 oy 8 = ? = oviduct = : ? as 4 5 .~ 3 ta) ie) rs = re oa »>?P enis, = £7 of i) = Ae fall eee ceptaculum seminis, v = vagina a Hs pr = prostate, 6 as & 5 e Q Qs ges g Es, , uscular vas de- Q SErECCEP RE USES s D o ae SP 7 Q have 24 1 we aS s amellae; they : o || = = D 2 are retractil ra a % so} a th : ile and are : =) Gl S| ese os ff ABS e anterior one-quarter of the body. They h Aer » OS eleh 8 8 56 & raised margin with tubercl ay aa y have a slightly TJ “e) =) Gal Gh! fads a seen call pock rcles similar to those on the dor = Sm en ctine pee ny MONE eee : pocket has a small, sli : iS) “4s 2) > 8 ToG Zu i , slightl i F = as Oe gs oS Olu Ww : g ral E = sa a, Mee the same-sized tubercles found a th aaa a rs Oo, Ss BFS & ody. It contains si Pits e rest of the = 2 a) S Six tripinnate gi : 5 5 a semi-circle around ihe post gills, three per side, set in = 2|s é P3 : erio 3 : pos] c Pas] . a. raised cone. r anus, which is on a S| sl|z 3 2 ae gee The ground colo : 5 eB) & & 2 g° 52 r varies from pale 7 S 2 3 ae > a g low to gold cream to bright yel a B=) - oO A 6B bl golden orange. There are dark b Sree. 2 wiles £2 4 Sag (0) 4 rown, (5) as} . tches, which are formed by cl speckled 9 3 = 3 Sm BE ing dark pigment at their b usters of tubercles hav: oF & iz e : ; ases and sid 6 =| light ti sides, but retain 3 eI BA the ps. These brown blotches are irregular, but | ae 5 5 8 E eo) central region, smaller on th ; arger S a2] oe 4 neous ue times they are composed of onl e mantle where some- S || e383] 2 a= E Ss £ $ & on i &§ oOo} © i) o Sr is There are also brown blotch yacue pigmented tubercle: , Ba) Z 2 Gl jo ease Has e 9 ° 3 TS ° : Been the body 5: s on the underside of the gO SA ees = £4. ixtacyslhores 1 y sides, and the top of the foot. Th a) = o s 4 25 s have cream b : € ases an Wt es orange lamellae; the gill d dark yellow or golden Q 2 & BS = O > gills are yellow or gold ass & Qs s n the ventral sid anf cee oa BS oe a Ommmene) ws ce e, which is the sam =| ea 3 = .8 Osan ground color, the poorly defi sae color as the Bao 88 42335 aie aah i ee y ned head is small and round = pclae 52 § a§5 a oS tacles on either sid pening. There are digitiform ten- 2|a 382 z : Ao ee ee side, 2 mm long, somewhat fl mDaows Sepa ea ho slightly wider at their bases. The foot 1 attened and yous Sg 2a5 588 32 2 oot is bilabi : ae| ira) oO pm) +o oH an notched on the upper lip. The ees ae = : 5 expanded anteri j is oval, not 5 RES eriorly, with a leaflike tail, and is wid e g = ung by the mantle widely Ble 3 é I v I eee we Yee Os nterna 9 : re q ternal anatomy: The body wall is 2-4 Bales AAS Z 8 spiculose, which alo : —4 mm thick and nN) 2 A a eal Os ng with the closely set spi = so S22 see aS) spiculose tu- 2 Sa 82 8 ® RG Oo AS) > 3 la s = g v The Veliger, Vol. 43, No. 4 Bis ee bs ee are SS Figure 4 A. Peltodoris mullineri, Millen & Bertsch, sp. nov. Specimen from San Clemente Island, November 1985. B. Notal tubercles. Scale bar = 400 zm. Paratype from Santa Cruz Island (CASIZ 74651). C. Radula. Scale bar = 1.5 mm. Specimen from San Clemente Island, November 1985. D. Small inner teeth. Scale bar = 100 um. Paratype from Santa Cruz Island (CASIZ 74651). E. Large medial teeth. Scale bar = 100 um. Paratype from Santa Cruz Island (CASIZ 74651). E Diminishing outer teeth. Scale bar = 40 um. Paratype from Santa Cruz Island (CASIZ 74651). S. V. Millen & H. Bertsch, 2000 hd Imm Figure 5 Reproductive system of Peltodoris mullineri, Millen & Bertsch, sp. nov. Scale bar = | mm. Specimen from Catalina Island (CASIZ 069302). Key: al = albumen gland, at = atrium, be = bursa copulatrix, ha = hermaphroditic ampulla, hd = hermaph- roditic duct, id = insemination duct, me = membrane gland, mu = mucus gland, od = oviduct, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = muscular vas deferens. bercles, gives the body its firm texture. The buccal tube is small in diameter and elongate with strong retractor muscles posteriorly. The buccal bulb is round with a long, broad, upwardly projecting radular sac emerging from the ventral posterior surface. Inside, the pale yellow lip disk is cuticular, thin, and smooth. The pale yellow radula has the formula 23-24 (42-63.0.42—63) (Figure 4C). The teeth are simple hooks, the central area has 12-15 small teeth per side (Figure 4D), which abruptly increase in size and remain constant (Figure 4E) until the last five to six which gradually diminish at the margin (Figure 4F). The salivary glands are long and ribbonlike with wide bases. The esophagus is long, wide, and tubular entering an elongate oval stomach, which is free of the digestive glands except at its posterior end. The digestive glands appear as one mass. No caecum was present. The flat intestine loops far to the right side and gradually narrows as it passes posteriorly to the anus. Page 361 The heart has a large triangular ventricle, a small elon- gate, diamond-shaped auricle, and blood vessels which end in two blood glands which are pink with black spots in preserved animals. The posterior gland is small, elon- gate oval, and located posterior to the central nervous system. The large anterior gland forms a triangle over the buccal mass. The central nervous system is granular and appears as one mass when viewed dorsally. On the ventral side, it can be seen that the small, oval cerebral ganglia are anterior and fused to the slightly narrower, granular pleural ganglia. The oval pedal ganglia are larger and more lateral. Under the salivary glands the small, round buccal ganglia lie close together. The reproductive system (Figure 5) is triaulic. The ovo- testis has several small, granular female acini peripheral to each oval male acinus. Clusters of stalked male acini coalesce into long ductules which empty into a common hermaphroditic duct. The narrow pre-ampullary duct wid- ens into a folded, tubular ampulla. Distally the ampulla bifurcates into a short oviduct and the prostatic portion of the vas deferens. The prostate is broad and folded; proximally it is white and smooth, distally it is cream colored and granular. The non-prostatic portion of the vas deferens is a thin, muscular, coiled tube which terminates in the common atrium forming a thick, muscular, un- armed penis. The elongate common atrium is internally plicate with a brown pigmented band near the exit. The vagina is a long, thin tube which exits the atrium and runs the length of the female gland mass to enter the large, spherical, thin-walled, orange bursa copulatrix. The insemination duct exits separately, in a serial arrangement, bending as it descends, and just before it enters the female gland mass, it is joined by the short stalk of the receptaculum seminis. The colorless receptaculum is pear-shaped and distally nodular. The female gland mass consists of a granular, yellow albumen gland, a large, folded, white mucus gland, and a smooth, creamy white membrane gland. The membrane gland extends into a distal, tubular, glandular nidamental duct which exits next to the common atrium. The repro- ductive openings appear as one non-protruding, plicate, depression located high on the night side, one-third of the way back from the anterior mantle margin. Natural history: This species has a known range from Santa Barbara and Anacapa Islands, California (Behrens, 1997) to Cabo San Lucas, Baja California, Mexico (Beh- rens, 1991). It has been found subtidally to 17 m in Au- gust, October, and December. It is presumed to feed on demosponges, but neither the food nor spawn mass have been identified. Peltodoris lancei Millen, sp. nov. (Figures 1C, 6A—E 7) Etymology: This species is named for Jim Lance, of San Diego, California, who was one of the first to know and love the opisthobranchs of the Gulf of California. Page 362 The Veliger, Vol. 43, No. 4 Figure 6 A. Peltodoris lancei Millen, sp. nov. Photo by Michael D. Miller. Paratype from Bahia de los Angeles (CASIZ 118573). B. Notal tubercles. Scale bar = 400 pm Paratype from Bahia de los Angeles (CASIZ 118572). C. Radula. Scale bar = | mm. Paratype from Bahia de los Angeles (CASIZ 118572). D. Small, inner teeth. Scale bar = 200 wm. Paratype from Bahia de los Angeles (CASIZ 118572). E. Large medial teeth. Scale bar = 100 pm. Paratype from Bahia de los Angeles (CASIZ 118572). E Diminishing outer teeth. Scale bar = 40 um. Paratype from Bahia de los Angeles (CASIZ 118572). S. V. Millen & H. Bertsch, 2000 Material examined: Holotype: CASIZ 110807, Punta la Gringa, Bahia de los Angeles, Gulf of California, Baja California, Mexico (29°02.56'N, 113°32.29'W), collected by Mike D. Miller, under rock, 7 m depth, on 28 June 1996, preserved length 55 mm, in Bouins solution. Par- atypes: CASIZ 118572, 1 specimen, dissected, found with the holotype, preserved length 58 mm, in alcohol. CASIZ 118573, Punta la Gringa, Bahia de los Angeles, | speci- men, 5 m depth, 30 June 1997, under rock, living length 27 mm, collected by Mike D. Miller. External morphology: This dorid nudibranch is large, with a living length of up to 75 mm. The largest specimen had a preserved size of 58 X 50 X 16 in a partially con- tracted state. The body shape is oval and depressed with a large overhanging crenulate mantle 15-20 mm wider than the foot. The notum is covered all over with closely set, conical papillae which end in white tips and are stiff- ened with spicules embedded in their sides (Figure 6B). The lamellate rhinophores are narrower in front and slope posteriorly; they have 15 fine lamellae on each side of their anterior edge and are strengthened with vertical spicules in the posterior half. They can each retract into a raised sheath, which on the sides has the same-sized papillae as on the notum, and slightly larger papillae at the edge. The gill pocket is almost smooth at the raised edge but has papillae on the sides which are the same as those on the notum. The branchial opening is wide and contains six to eight tripinnate gills, which are arranged three or four per side around the posterior anus, which is on a raised cone. The holotype has two small pinnate branches on the anterior midline, which appear to be ex- tensions from the anteriormost gill on each side. The ground color is dark orange-brown to reddish or- ange. The top one-fourth of each papillae is opaque white giving a uniform speckling of tiny spots evenly distrib- uted over the notum. Toward the mantle edge, the tuber- cles, and hence the spots, become smaller, and the ground color lightens to form a bright yellowish orange rim to the mantle. The smallest, darkest specimen had a few faint brownish spots, one in front of the rhinophores, two on each side between the rhinophores and gills, and one on the right side posterior to the gills (Figure 6A). The rhinophores and gills are dusky yellow with lighter yel- low tips. On the underside, the mantle and foot are yel- low. The tops of both the head and foot are orange. Ventrally, the head is oval, hard, and granular on top with a vertical mouth and long digitiform tentacles on each side, reaching 3—3.5 mm. The sides of the body are short, up to 3 mm high, and the underside of the mantle shows light radial striations indicating the presence of spicules. The foot is elongate oval with a slightly tapered tail, which is overhung by the mantle all around. It is hard and granular dorsally; the anterior edge is deeply bilabiate and notched. Internal anatomy: The thick body wall contains a mesh- Page 363 work of bundles of fine spicules and connective tissue. The buccal tube is short and muscular with strong retrac- tor muscles. The buccal bulb is round and short with a round, ventral, slightly projecting radular sac. Inside, the lip disk is round, smooth, and colorless. The pale yellow radula has the formula 28(33.0.33). The rachis area is wide and bare (Figure 6C). The teeth are simple hooks, with no denticulations. There are four small teeth near the bare rachis, (Figure 6D) then they increase rapidly in size and are uniform (Figure 6E) except the four outer- most teeth on each side which rapidly decrease in size (Figure 6F). The salivary glands are white and dense, forming wide straps on either side of the buccal bulb, narrowing as they go through the nerve ring, and becom- ing long straps bent back on themselves. The esophagus is a thin-walled wide tube, which at the notch of the di- gestive glands becomes a saclike stomach situated in a hollow between the otherwise confluent digestive glands. The intestine is wide and muscular where it leaves the stomach and then it loops to the right and across the top of the digestive glands to end on a small cone at the posterior edge of the gills. The heart is large, ending in two leaf-shaped, yellow blood glands. The smaller anterior gland is slightly an- terior to the central nervous system; the larger posterior gland is over the esophagus. The central nervous system is enclosed in a tough semi-transparent sheath and is smooth dorsally, granular ventrally. The elongate oval cerebro-pleural ganglia are fused; the smaller, oval pedal ganglia are posterior and slightly ventral. The common commissure is enclosed in a thick sheath. There is a small, unpaired visceral ganglion. The buccal ganglia are very small and a short distance apart, each with a small gastroesophageal ganglion connected by a short nerve. The reproductive system (Figure 7) is triaulic. The yel- low gonad covers all sides of the digestive glands. The ovotestis has round acini leading to the hermaphroditic duct, which leaves the digestive glands to the right of the esophagus. The hermaphroditic ampulla is a long, grey, sinuous tube, which narrows abruptly, dividing into a short oviduct which enters the base of the albumen gland and the vas deferens. After a short distance, the vas de- ferens widens into a massive, U-shaped prostate gland. The proximal portion is creamy white and finely granular; the distal one-third is yellow and coarsely granular. Fol- lowing the prostatic portion, the vas deferens is a long, narrow threadlike tube, which folds a few times, then runs straight down the outer surface of the female gland mass to the penis. The penis is small, muscular, and bulbous with a pointed, colorless, lightly cuticular tip. The vagina is posterior and separate from the male opening. Inter- nally it is soft and plicate at the atrium, then villous for a short distance before narrowing. The tubular vagina en- ters a large, irregular oval bursa copulatrix. In its normal position it is almost completely hidden by the prostate gland. The insemination duct leaves the bursa near the Page 364 The Veliger, Vol. 43, No. 4 vd be mu rs Figure 7 Reproductive system of Peltodoris lancei Millen, sp. nov. Par- atype from Bahia de los Angeles (CASIZ 118572). Scale bar = 1 mm. Key: al = albumen gland, be = bursa copulatrix, ha = hermaphroditic ampulla, hd = hermaphroditic duct, id = insem- ination duct, mu = mucus gland, od = oviduct, p = penis, pr = prostate, rs = receptaculum seminis, v = vagina, vd = muscular vas deferens. vaginal entrance in a serial arrangement. Just before it enters the base of the albumen gland it is joined by a short duct from the almost sessile, elongate-oval recep- taculum seminis. The female gland mass is hard and com- pact. The largest portion, the mucus gland, is dorsal and pink and cream in color. It wraps around the small, oval, yellowish albumen gland. The distal membrane gland is soft and white, wrapped around the separate nidamental opening. The genital openings are located approximately one-third of the way back from the anterior edge of the mantle. The penis opening is anterior with a posterior crescent-shaped slit leading to the vaginal opening. The ventral nidamental opening is separated by a small papilla and a ridge of tissue. Natural history: This species is known only from Punta la Gringa, Bahia de los Angeles, Baja California, Mexico. A pair was found in June, both large and apparently ma- ture; one smaller specimen was found in June the follow- ing year. No spawn was seen. The animals were feeding on a yellow encrusting demosponge on the undersides of rocks. Discussion of Peltodoris: The two new species described here have the following characteristics of the genus Pel- todoris. They have depressed, rigid bodies with a wide margin, and minute, small, evenly disposed spiculose tu- bercles and a small head with digitiform tentacles. Inter- nally they have a smooth lip cuticle, simply hooked rad- ular teeth with smaller teeth toward the center of the rad- ula abruptly increasing in size, elongate salivary glands, and a stomach free of the digestive glands. The repro- ductive system has a serially arranged bursa copulatrix, an enlarged prostate gland, and an unarmed penis. There are 14 described species of Peltodoris, of which all but five can be separated from the two species de- scribed here because they do not have a yellow or orange ground color. Those five are compared with the two new- ly described species in Table 2, and differences are point- ed out in the discussion below. Peltodoris angulata Eliot, 1904, can be separated because it has three violet-brown spots on each side between the rhinophores and gills and a peculiar foot which is anteriorly extended into tentac- ular angles 3.5 mm long. Peltodoris aurea Eliot, 1904, has dull violet spots along the mantle edge and a spot in front of the gills and one behind the rhinophores. The tubercles are in the form of small, flat warts. The color photograph by Orr (1981:44) does not appear to fit the original description of this species. Peltodoris nayarita Ortea & Llera, 1981 is similiar to P. mullineri sp. nov., but has pinnate brown gills, brown rhinophores, and smaller spots on the body. Peltodoris noumeae Risbec, 1937, is orange-yellow and has white-tipped tubercles like P. lancei, but also has irregular bright, “‘fiery”’ spots, which are not found in the latter species. Peltodoris punc- tifera (Abraham, 1877) has white-tipped tubercles like those of P. lancei, but it also has small violet-brown or grey spots on the notum and brown speckles on the rhin- ophores and gills. In southern California and on the Pacific coast of Mex- ico, Peltodoris mullineri sp. nov., is most easily confused with P. nayarita and Jorunna pardus Behrens & Hen- derson, 1981. It can be separated from both of these spe- cies because it lacks dark brown on the gills and rhin- ophores. In the Gulf of California, Peltodoris lancei is easily confused with the porostome nudibranchs Doriop- silla albopunctata (Cooper, 1863) and Doriopsilla gemela (Gosliner et al., 1999). It can be distinguished externally by its firm texture, smaller and more uniform white spots, always located on conical spiculose tubercles, and con- tinuing to the mantle edge, and the presence of long dig- itiform tentacles on the head. Acknowledgments. We thank Michael D. Miller for finding and photographing Peltodoris lancei (Figures 1C, 6A). We are also grateful to Jeff Hamann for providing us with photographs 1A and B and funding for the color plate. Terry Gosliner and the California Academy of Sciences provided us with several spec- imens, collecting data, and assistance with SEM radula photo- graphs 2F and 4C. This research was partly funded by the De- S. V. Millen & H. Bertsch, 2000 Page 365 Table 2 Peltodoris species with yellow or orange ground color. Rhinophore Bursa Species Ground color Tubercles color Gills Radula copulatrix angulata Eliot, yellow? (pres.), 3 minutely granular ? 6 tripinnate 38 (45.0.45) ? 1904 violet-brown spots each side aurea Eliot, sand, dull violet small flat, orange warts ? 8 tripinnate 20 (25.0.25) y 1904 spots on edges, 2 on midline lancei sp. nov. red-orange fine conical spiculose red-orange 6-8 tripinnate, 28 (33.0.33) serial tubercles, white- red-orange tipped mullineri sp. light yellow, brown granular, spiculose tu- golden yellow 6 tripinnate, 23-24 (42— serial nov. blotches bercles golden yel- 63.0.42—-63) low nayarita Ortea yellow-orange, fine conical spiculose brown 8 unipinnate, 51 (50.0.50) semi-serial & Llera, brown flecks tubercles brown 1981 noumeae Ris- yellow-orange, spiculose, white-tipped yellow, white 4 bipinnate, 2 ? (20—35.0.20—35) — serial becwl937 bright fiery spots ups pinnate, yel- low, white spots punctifera yellow, large brown fine conical tubercles, white & brown’ 6 bi- & tripin- 30 (50.0.50) serial (Abraham, or grey spots dark specked, largest nate, dark 1877) white-tipped spots, white tips partment of Zoology, University of British Columbia. We are grateful to the Mexican federal government agency Comision Nacional para el Conocimiento y el Uso de la Biodiversidad (CONABIO) for the grant which enabled HB to perform research at Bahia Tortugas; we especially thank Carlos Sanchez, José Luis Arreola, and Orso Angulo (Universidad Aut6noma de Baja Cal- ifornia Sur, La Paz); and Manuel Higuera Serrano (Cooperativa La Purisima) and Francisco Javier Zacatzi Ayala (both from Ba- hia Tortugas) for assistance in the field. LITERATURE CITED BEHRENS, D. W. 1980. Pacific Coast Nudibranchs, A Guide to the Opisthobranchs of the Northeastern Pacific. Sea Chal- lengers: Los Osos, California. 112 pp. BEHRENS, D. W. 1983. Report of the 1982 outer Baja California expedition. Opisthobranch Newsletter 15(4):18—-19. BEHRENS, D. W. 1991. Pacific Coast Nudibranchs. 2nd ed. Sea Challengers: Monterey, California. vi + 107 pp. BEHRENS, D. W. 1996. Peltodoris sp. http://siolibrary.ucsd.edu/ slugsite/nudiwkps/nudiwk27.html BEHRENS, D. W. 1997. Range extensions of eight northeastern Pacific nudibranchs. Opisthobranch Newsletter 23(4):13. BEHRENS, D. W. & R. HENDERSON. 1981. Two new cryptobranch dorid nudibranchs from California. The Veliger 24(2):120— 128. BertTscH, H. 1989. Phylum Mollusca: Subclass Opisthobranchia (shell-less snails, nudibranchs) Pp. 54—68 in A. Kerstitch, Sea of Cortez Marine Invertebrates. Sea Challengers: Mon- terey, California. BertscuH, H. 1991a. Seasonal, geographic and bathymetric dis- tribution of opisthobranchs at Bahia de los Angeles, Baja California, México. Western Society of Malacologists, An- nual Report 23:18. BerTSCH, H. 1991b. Biodiversity of the dorid nudibranch (Mol- lusca: Opisthobranchia) fauna of the Pacific coast of the three Californias: systematic and zoogeographic comments. Pp. 107-115 in M. Dailey & H. Bertsch (eds.), Proceedings of the VIII International Symposium of Marine Biology, En- senada, Baja California, México. BerTScH, H. 1995a. A three-year phenological analysis of sub- tidal populations of opisthobranch molluscs at Bahia de los Angeles, Baja California, Mexico. Proceedings of the Pacific Division of the American Association for the Advancement of Science 14(1):34. BERTSCH, H. 1995b. Biogeographic microcosms: ecological con- trasts and variations in long-term regional faunal studies (Gulf of California, Mexico). Western Society of Malacol- ogists Annual Report 27:4—5. BeRTSCH, H. 1996. Trapania slugsite/nudiwk38.html. BerTScH, H. & A. KersTITCH. 1984. Distribution and radular morphology of various nudibranchs (Gastropoda: Opistho- branchia) from the Gulf of California, Mexico. The Veliger 26(4):264—273. CERVERA, J. L. & J. C. GARCIA-GOMEZ. 1989. Dos nuevas espe- dies de Trapania Pruvot-Fol, 1931 (Gastropoda: Nudibran- chia) del sur de espana. Bolletino Malacologico 24:189—204. GosLinER, T. M. 1987. Nudibranchs of Southern Africa, a Guide to Opisthobranch Molluscs of Southern Africa. Sea Chal- lengers: Monterey, California. 136 pp. GOSLINER, T. M. & D. W. BEHRENS. 1986. Two new species and genera of aeolid nudibranchs from the tropical eastern Pa- cific. The Veliger 29(1):101—113. sp. http://siolibrary.ucsd.edu/ Page 366 GOSsLINER, T. M., M. T. GHISELIN & H. BERTSCH. 1985. Opistho- branch mollusks of the Punta Eugenia region, with a dis- cussion of biogeographical affinities. Western Society of Malacologists Annual Report 17:11. GOSLINER, T. M., M. C. SCHAEFER & S. V. MILLEN. 1999. A new species of Doriopsilla (Nudibranchia: Dendrodorididae) from the Pacific coast of North America, including a com- parison with Doriopsilla albopunctata (Cooper, 1863). The Veliger 42(2):181—190. KEEN, A. M. 1971. Sea Shells of Tropical West America; Marine Mollusks from Baja California to Peru. Stanford University Press: Stanford, California. xiv + 1064 pp. Lance, J. R. 1983. California underwater. California Magazine 8(3):85-89. The Veliger, Vol. 43, No. 4 Orr, J. 1981. Hong Kong Nudibranchs. Urban Council: Hong Kong. 82 pp. PIcTON B. & C. C. Morrrow. 1994. A Field Guide to the Nu- dibranchs of the British Isles. Immel Publishing: London. 143 pp. ROLLER, R. A. & S. J. LONG. 1969. An annotated list of opistho- branchs from San Luis Obispo County, California. The Ve- liger 11(4):424—430. RupMAN, W. B. 1987. The genus Trapania (Nudibranchia: Gon- iodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53:189—212. SKOGLUND C. 1991. Additions to the Panamic Province Opis- thobranchia (Mollusca) literature 1971-1990. The Festivus 22 (Suppl. 1):ii+27 pp. The Veliger 43(4):367—375 (October 2, 2000) THE VELIGER CMS, Inc., 2000 A New Species of Hydrobiid Snail of the Genus Pyrgulopsis from Northwestern Nevada ROBERT HERSHLER Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0118, USA AND DONALD W. SADA Desert Research Institute, 2215 Raggio Parkway, Reno, Nevada 89512 and Biological Resources Center, University of Nevada, Reno, Nevada 89557, USA Abstract. Pyrgulopsis bruesi new species, from northwest Nevada, differs from closely similar P. augustae on the basis of its higher-spired shell, larger penial filament, and more ventrally positioned seminal receptacle. This species is endemic to a thermal spring area in a small basin which was inundated by pluvial Lake Lahontan. INTRODUCTION Pyrgulopsis is the largest genus of freshwater mollusks in North America, with more than 130 species currently placed in this group (Hershler, 1994, 1995; Thompson, 1995; Hershler, 1998). Whereas these small, gill-breath- ing snails are broadly distributed in southern North Amer- ica, the greatest diversity of Pyrgulopsis is in the Great Basin, a vast expanse of internal drainage lying between the Wasatch Front and Sierra Nevada of the western Unit- ed States. Within this region most congeners are restricted to seeps and spring sources, while only a few species live in more integrated stream habitats. The isolated nature of these water bodies, reinforced by surrounding arid envi- ronments, has been highly conducive to differentiation of these poorly dispersing snails. Eighty species of Pyrgu- lopsis are now known from the Great Basin (Hershler & Sada, in press). We recently collected biota at Fly Ranch Hot Springs (also known as Ward’s Hot Springs; Garside & Schilling, 1979), north of Gerlach, Nevada. The small invertebrate assemblage of mollusks and crustaceans inhabiting ther- mal waters at this locality, which is part of a well-defined northeast-trending zone of thermal springs in northwest Nevada (Hose & Taylor, 1974:fig. 2), included a new spe- cies of Pyrgulopsis, which is described below. This new species is most similar morphologically to several con- geners also living in the Great Basin of Nevada. MATERIALS anD METHODS Specimens are deposited in the National Museum of Nat- ural History, Smithsonian Institution (USNM). Terminol- ogy and methods of morphological analysis are of Her- shler (1998) and Hershler & Ponder (1998). Measure- ments of shells of the holotype and a series of paratypes are in Table 1. SYSTEMATICS Pyrgulopsis bruesi Hershler & Sada, sp. nov. Type material: The holotype (USNM 892079) is a dried shell (2.55 mm shell length, Figure 1) from a small stream which enters Fly Reservoir, about 23 airline-km north-northeast of Gerlach, Washoe County, Nevada, T. 34 .N., R. 23 E., SE % section 2, elevation about 1238 m; collected by R. Hershler and D. W. Sada, 19 July 1997. Paratypes (USNM 860868) consist of a large series of dry shells and alcohol-preserved specimens collected from the type locality at the same time. Two additional series (USNM 892075, 892584), which are not designated as paratypes, were collected at the type locality by D. W. Sada at other times in 1997. Diagnosis: A medium-sized species with ovate-conic shell having weakly rounded whorls. Penis small, narrow bladelike; filament medium length, lobe and penial glands absent. Description: Shell (Figure 2A—C) ovate-conic, almost pupiform, width/height, 49-58%; height, 2.02—3.03 mm; width, 1.13—1.74 mm; whorls, 4.5—5.25. Protoconch near planispiral, 1.4 whorls, diameter about 300 wm, earliest 0.5 whorl slightly wrinkled, otherwise smooth (Figure 2D-F). Teleoconch whorls weakly convex or nearly flat (earlier whorls somewhat more rounded), sometimes nar- rowly shouldered, basal portion sometimes strongly an- Page 368 Table | Morphometric and meristic shell features of holotype and ranges of values for 10 paratypes of Pyrgulopsis bruesi, new species. Morphometric parameters are expressed in mm. Holotype Paratypes Shell height 2.55 2.38—2.75 Shell width 1.47 1.19-1.47 Body whorl height 1.82 1.56—1.90 Body whorl width 127) 1.15-1.41 Aperture height 0.99 0.85—1.07 Aperture width 0.87 0.82—0.93 Shell width/shell height 0.57 0.49-0.58 Number of whorls 4.75 4.75-5.25 gled; sometimes loosened for 0.25 whorl behind aperture. Aperture ovate, broadly adnate or slightly disjunct. Inner lip complete, slightly thickened; columellar lip sometimes slightly reflected. Outer lip thin, orthocline or weakly pro- socline, sometimes weakly sinuate. Umbilicus narrow. Periostracum tan. Shells of males narrower, with flatter whorls, and slightly smaller than those of females. Operculum (Figure 2G—I) thin, light amber throughout or colored only in attachment region, ovate, multispiral, nucleus eccentric. Edges of whorls frilled on outer sur- face (Figure 2G, H), sometimes markedly so; outer mar- gin sometimes having weak rim. Attachment scar margin strongly thickened between nucleus and inner edge (Fig- ure 21), sometimes thickened all around. Buccal mass medium-sized; radular sac extending be- hind buccal mass as small loop. Radula about 530 x 80 pm, with 55—60 rows of teeth. Central teeth (Figure 3A, B) trapezoidal, about 18.6 1m wide, dorsal edge well in- dented; lateral cusps 4—6; central cusp medium width, proximal edges parallel, distal portion pointed; basal cusp 1, large. Basal tongue broad V-shaped, basal sockets deep. Lateral teeth (Figure 3C) with 2—3 cusps on inner side and 3-5 cusps on outer side, central cusp rather broad; neck weakly flexed; outer wing length 200% width of tooth face. Inner marginal teeth (Figure 3D) with 21— 27 cusps, outer marginal teeth (Figure 3E, F) with 23-27 cusps. Style sac about as long as remainder of stomach; stomach with small posterior caecum. Cephalic tentacles usually unpigmented, sometimes pigmented with a few black granules proximally. Snout The Veliger, Vol. 43, No. 4 Figure 1 Holotype of Pyrgulopsis bruesi Hershler & Sada, sp. nov., from Fly Hot Springs, Washoe County. Nevada. unpigmented. Foot usually unpigmented, sometimes pig- mented with scattered black granules dorsally. Opercular lobe black along inner edge. Neck variably pigmented with black granules. Pallial roof, visceral coil lightly pig- mented to near black, pigment lighter on genital duct than elsewhere. Males usually with darker pigment on mantle roof and visceral coil than females. Filament of penis al- most uniformly pigmented with internal black granules, granules scattered proximally along length of penial duct. Dorsal and ventral surfaces of tentacles bearing single longitudinal ciliary tracts. Distal penis bearing scattered long cilia near terminus, otherwise unciliated. Ctenidium well developed; filaments 18-20, with pleats, broadly triangular, apices centrally positioned; cte- nidium abutting pericardium posteriorly. Osphradium rather long, intermediate width, near centrally positioned. Renal gland longitudinal; kidney with small pallial bulge, opening slightly thickened. Rectum near straight, broadly overlapping pallial oviduct, abutting prostate gland. Ovary 0.75 whorl, filling less than 50% of digestive gland behind stomach, abutting posterior edge of stom- ach. Albumen gland (Ag) having substantial pallial com- Figure 2 Scanning electron micrographs of shell and opercula of P. bruesi Hershler & Sada, sp. nov. USNM 860868. A-C. Variation in shell shape (shell height, 2.58 mm, 2.43 mm, 2.68 mm, respectively). D—F Shell apices, showing weak protoconch sculpture (bars = 120 wm). G, H. Outer surface of operculum (bars = 240 ym, 200 pm, respectively). I. Inner surface of operculum, showing thickened border of muscle attachment area near nucleus (bar = 240 ym). R. Hershler & D. W. Sada, 2000 Page 369 The Veliger, Vol. 43, No. 4 Figure 3 Scanning electron micrographs of radula of P. bruesi Hershler & Sada, sp. nov. USNM 860868. A, B. Central radular teeth (bars = 9.2 wm, 10.9 wm, respectively). C. Lateral tooth (bar = 10.9 pm). D. Inner marginal tooth (bar = 12.0 pm). E, EF Outer marginal teeth (bars = 12.0 wm). ponent (Figure 4A). Capsule gland (Cg) slightly shorter, about as wide as albumen gland, divided into two equal- sized tissue sections (anterior, clear; posterior, white), broadly ovate in section, right lobe thicker than left; rectal furrow absent. Ventral channel (Vc) broadly overlapping capsule gland; longitudinal fold well developed. Genital aperture (Ga) a long terminal slit. Coiled oviduct (Cov) of two overlapping posterior-oblique loops. Oviduct and bursal duct joining just behind pallial wall. Bursa copu- latrix (Bu) small relative to albumen gland, narrow or ovate, longitudinal, up to 50% of length posterior to al- bumen gland. Bursal duct originating from anterior edge at midline, slightly longer than bursa copulatrix, narrow or medium width (Figure 4B). Seminal receptacle (Sr) small relative to bursa copulatrix, ovate, positioned along ventral edge of distal oviduct coil and usually just anterior R. Hershler & D. W. Sada, 2000 Page 371 Bu Cov Figure 4 Genitalia of P. bruesi Hershler & Sada, sp. nov. USNM 860868. A. Left side of female glandular oviduct and associated structures. B. Bursa copulatrix and its duct. C. Seminal receptacle and its duct. D. Left side of prostate gland, showing insertion of vas deferens. E. Penis. Bars = 0.25 mm. Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cov = coiled oviduct, Ga = genital aperture, Pd = penial duct, Pvd = pallial vas deferens, Pr = prostate gland, Pw = posterior wall of pallial cavity, Sr = seminal receptacle, Vc = ventral channel of capsule gland, Vvd = visceral vas deferens. to bursa copulatrix, duct slightly shorter than body (Fig- ure 4C). Testis 1.5—2.0 whorls, composed of numerous com- pound lobes, filling about 50% of digestive gland behind stomach, broadly overlapping stomach anteriorly. Semi- nal vesicle a medium-sized mass of tight coils opening from and positioned alongside the anterior portion of tes- tis. Prostate gland (Pr) small, bean-shaped, pallial portion about 33% of total length, narrowly ovate in section (Fig- ure 4D). Visceral vas deferens (Vvd) opening to ventral edge of prostate gland just behind pallial wall. Pallial vas deferens (Pvd) opening to ventral edge of prostate gland just in front of posterior wall; duct having distinct prox- imal bend on columellar muscle. Penis small; base elon- gate-rectangular, smooth; filament about 33% length of base, gently tapering, distally pointed, rather broad, lon- gitudinal, poorly distinguished from base (apart from darker pigmentation); lobe and glands absent (Figure 4E). Penial duct (Pd) near inner edge, narrow, undulating a few times near base, otherwise near straight. Etymology: A patronym, named after the late Charles Thomas Brues II (1879-1955) who, in addition to his notable studies in entomology (Mallis, 1971), surveyed Page 372 thermal springs of the western United States (Brues, 1928, 1932) and brought the interesting biota of these habitats to the attention of the biological community. Brues (1932:206) early visited the type locality area (‘‘sixteen miles northwest of Gerlach’’), although he did not collect any snails. We propose the vernacular name, “Fly Ranch pyrg,”’ for this species in reference to its type locality. Comparisons: Pyrgulopsis bruesi and a small group of congeners share a simple penis without lobe or glands. Among these, P. bruesi is most similar to P. augustae Hershler, 1998, which lives about 190 km to the southeast (Antelope Valley, Lander County), but differs on the basis of its higher-spired shell (compare Figure 2A—C with Hershler, 1998:fig. 9B), larger penial filament (compare Figure 4E with Hershler, 1998:fig. 40A), and more ven- trally positioned seminal receptacle (compare Figure 4A with Hershler, 1998:fig. 40B). These two species, together with another congener from the Great Basin of Nevada (P. dixensis Hershler, 1998) having a simple penis, are united by their narrow shell and distinctive thickening of the operculum attachment region border near the nucleus. Habitat: The Fly Ranch Hot Springs is the largest spring zone in northwest Nevada (Garside & Schilling, 1979), consisting of about 40 sources (Grose & Keller, 1975) that collectively discharge about 500 acre-feet/year (= 0.69 cfs) (Sinclair, 1962). Thermal sources are located on a large mound and discharge into a series of pools, with drainage ultimately entering the Fly Reservoir (Figure 5). This hot spring area has been modified by surface water diversion, groundwater mining, dredging, and recreation- al activity. Pyrgulopsis bruesi was collected from a turbid, 1 m deep and 1.5 m wide stream (Figure 6) fed by natural springs and a flowing well whose effluents have given rise to an impressive, three-pronged travertine tower (a second, similarly active tower, slightly north of the above, was figured by Sinclair, 1962:fig. 1; and Garside & Schil- ling, 1979:74). Water quality data obtained from this stream on 20 September 1997 are as follows: water tem- perature, 27.0°C; conductivity, 1760 mhos/cm; pH; 8.2; and dissolved oxygen content, 6.7. Snails were collected from submerged unidentified aquatic vegetation on which they were moderately abundant; fewer specimens were found on Cattail (Typha). Although considerably ceoled relative to the boiling effluent of the well, this stream nonetheless ranks as a regional thermal water body as variously defined (e.g., Waring, 1965:4; Garside & Schil- ling, 1979:1). Another hydrobiid snail, Tryonia protea, which is broadly distributed in western North America (Hershler, 1999); pulmonate gastropods, Helisoma anceps and Physella virgata, and an unidentified amphipod were collected in association with P. bruesi. The tui chub, Gila bicolor, also is present in this stream (Brues, 1932). The Veliger, Vol. 43, No. 4 Distribution: This species is thus far known only from its type locality. During the course of our field survey, other sites in the Hualapai Flat basin were searched and did not yield this snail, although a congener, P. gibba, was collected from the flanking Granite Range (Hershler, 1998). DISCUSSION Pyrgulopsis has occupied habitats of the northern Great Basin since at least the late Miocene (Hershler & Sada, in press), and several scenarios for the origin of P. bruesi may be conjectured. Hualapai Flat is a small (about 200 km/?), endorheic, north-south trending graben bounded to the west by the imposing Granite Range. The Calico Hills form a lower hydrographic barrier to the northeast, while to the southeast Hualapai Flat is separated from the Black Rock Desert by an alluvial divide of only about 30 m (Sinclair, 1962). Thick Quaternary deposits on the floor of Hualapai Flat include fine sections attributable to Lake Lahontan, which also left sedimentary exposures else- where in the basin (Grose & Sperandio, 1978). Highstand elevations of Lake Lahontan (from Mifflin & Wheat, 1971) imply that the Fly Hot Springs area was flooded by a > 100 m deep embayment during the late Pleisto- cene. Evolution of P. bruesi may be attributable to iso- lation in basinal spring habitats coincident with the final recession of Lake Lahontan about 13,000 ybp. Pyrgulopsis bruesi is one of many congeners that are endemic to flowing thermal waters. Experimental data in- dicates that while these snails actively prefer thermal hab- itats, they can tolerate considerably lower water temper- atures (Mladenka, 1992). Invasion of Hualapai Flat from elsewhere in the Lahontan basin via dispersal within plu- vial waters may thus be tenable for progenitors of P. bruesi. Congeners that are most similar morphologically to P. bruesi also live within (P. augustae) or adjacent to (P. dixensis) the Lahontan basin, supporting a hypothesis that these were all derived from a widespread pluvial pro- genitor. Note that recently acquired evidence of high el- evation early Pleistocene shorelines (about 1400 m) in the Lahontan basin implies that Dixie Valley (which harbors P. dixensis) may have been integrated with this regional drainage (Reheis, 1996; Reheis & Morrison, 1997), in- stead of containing an isolated pluvial lake as tradition- ally viewed (Mifflin & Wheat, 1979). Alternatively, one may attribute less importance to plu- vial dispersal of snails and postulate an earlier (pre-Pleis- tocene) origin of P. bruesi coincident with regional de- velopment of the modern basin and range landscape. To- pographic closure of Hualapi Flat during the Neogene may be inferred by its location along the edge of the Black Rock-Carson Sink zone of extension, a tectonically active area of broad alluviated grabens whose youthful development involved extensive deformation of 17—6 ma extrusive igneous rocks (Wallace, 1984). On a local scale, Page 373 R. Hershler & D. W. Sada, 2000 location NEVADA /W, MV aww iS Figure 5 Location of type locality of P. bruesi Hershler & Sada, sp. nov. in Hualapai Flat (arrows). Map from USGS Hualapai 7.5 minute series (topographic). Foothills of the Granite Range are to the west, and the edge of the Hualapai playa is to the southeast. > Flat South Quadrangle northwest-trending fault zone, which permits deep pene- Hualapai Flat has been tectonically active from the late tration to a regional heat reservoir (Sperandio & Grose, 1976). (Near-surface intrusive bodies are not sufficiently Cenozoic through the Holocene, with both normal and > lateral slip along several major fault zones (Grose, 1978; Grose & Sperandio, 1978). Uplift of the Granite Range on the west side of the basin has been particularly rapid and extensive (Bonham, 1969; Grose, 1978). Hualapai Flat is bisected by a regional north-south fault zone which extends from the High Rock area to the Pyramid Lake basin (Grose, 1987). Occurrence of local hot spring ac- hot to generate local geothermal activity.) Although in- dividual springs in the Hualapai Flat may be no more than Lahontan in age, the supplying heat reservoir has been in place for 2—20 my (Grose, 1978) which, together with locally extensive faulting from mid-Tertiary into the Ho- locene, suggests that a long history of thermal spring ac- feasible. tivity in the area now occupied by this basin is T, tivity is attributed to the intersection of this and anothe Page 374 Figure 6 Type locality of P. bruesi Hershler & Sada, sp. nov. Granite Range is in the background. Photograph, 19 July 1997. A pre-Pleistocene origin of P. bruesi requires persis- tence of this snail through the time period when Lake Lahontan flooded Hualapai Flat. Snails may have invaded basinal habitats from higher elevation springs (following desiccation of this lake) in conjunction with migrating geothermal activity as mediated by local faulting. (Fossil spring deposits attest to local migration of surface thermal activity; Grose, 1978.) The possibility that basinal springs harbored snails while covered by Lake Lahontan also merits consideration. Acknowledgments. We thank Y. Villacampa for assistance with scanning electron microscopy, K. Darrow for inking anatomical drawings, M. Ryan for her drawing of the holotype and assembly of plates, and D. Cole for preparing the map (all from USNM),. Fieldwork was supported, in part, by a 1997 award from the National Museum of Natural History Biological Surveys and In- ventories Program (BSI). LITERATURE CITED BONHAM, H. F. 1969. Geology and mineral deposits of Washoe and Storey Counties, Nevada. Nevada Bureau of Mines and Geology Bulletin 70:1—140, pls. 1-6. The Veliger, Vol. 43, No. 4 Bruges, C. T. 1928. Studies on the fauna of hot springs in the western United States and the biology of thermophilous an- imals. 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[Unpublished] Progress report [to the National Science Foundation] No. 4, for period February 1—October 31, 1975. 109 pp. Grose, L. T. & R. J. SPERANDIO. 1978. Geology of the Gerlach- Hualapai Flat geothermal area, northwestern Nevada. Pp. 1— 10 in G. V. Keller & L. T. Grose (eds.), Studies of a Geo- thermal System in Northwestern Nevada, Part 1. Colorado School of Mines Quarterly 73(3). HERSHLER, R. 1994. A review of the North American freshwater snail genus Pyrgulopsis (Hyrdobiidae). Smithsonian Contri- butions to Zoology 554:1—115. HERSHLER, R. 1995. New freshwater snails of the genus Pyrgu- lopsis (Rissooidea: Hydrobiidae) from California. The Ve- liger 38(4):343-373. HERSHLER, R. 1998. A systematic review of the hydrobiid snails (Gastropoda: Rissooidea) of the Great Basin, western United States. Part I. Genus Pyrgulopsis. The Veliger 41(1):1—132. HERSHLER, R. 1999. A systematic review of the hydrobiid snails (Gastropoda: Rissooidea) of the Great Basin, western United States. 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C. 1992. The ecological life history of the Bru- neau Hot Springs Snail (Pyrgulopsis bruneauensis). Final Report to the United States Fish and Wildlife Service. 116 pages. R. Hershler & D. W. Sada, 2000 REHEIS, M. C. 1996. Old, very high pluvial lake levels in the Lahontan basin, Nevada: evidence from the Walker Lake basin. United States Geological Survey Open-File Report 96-5 14:1-19. REHEIS, M. C. & R. Morrison. 1997. High, old pluvial lakes of western Nevada. Pp. 459—492 in P. K. Link & B. J. Kowallis (eds.), Proterozoic to Recent Stratigraphy, Tectonics, and Volcanology, Utah, Nevada, Southern Idaho and Central Mexico. Brigham Young University Geology Studies 42(1). SINCLAIR, W. C. 1962. Ground-water resources of Hualapai Flat, Washee, Pershing, and Humboldt Counties. State of Nevada Department of Conservation and Natural Resources Ground- water Resources—Reconnaissance Series Report 11:1—15, figs. 1-5, tables 1—4. SPERANDIO, R. J. & L. T. Grose. 1976. Tectonic controls on the Page 375 Fly Ranch hot spring system, Hualapai Flat, northwest Ne- vada. Geological Society of America Annual Meetings Ab- stracts with Programs 8(6):1116. THompson, F G. 1995. A new freshwater snail from the Coosa River, Alabama (Gastropoda: Prosobranchia: Hydrobiidae). Proceedings of the Biological Society of Washington 108(3): 502-507. WALLACE, R. E. 1984. Patterns and timing of late Quaternary faulting in the Great Basin Province and relation to some regional tectonic features. Journal of Geophysical Research 89(B7):5763—-5769. WarRING, G. A. 1965. Thermal springs of the United States and other countries of the world—a summary. United States Geological Survey Professional Paper 492:1—383. [revised by R. R. Blankenship & R. Bentall] The Veliger 43(4):376 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 NOTES, INFORMATION & NEWS Stohler Memorial Fund The Board of Directors of The Veliger met recently and discussed options for honoring Dr. Rudolf Stohler, The Veliger’s Founding Editor. It was the Board’s decision to create a memorial fund and invite members and friends to make contributions in Dr. Stohler’s memory. Donations to the Stohler Memorial Fund will be used to enhance the endowment that provides grants to students with lim- ited resources to cover their page charges. As Dr. Stohler provided many young malacologists with their first op- portunity to publish in a scientific journal in the pages of The Veliger, this seems a most fitting tribute to his mem- ory. Tax-deductible contributions may be directed to: The Veliger—Stohler Memorial Fund Dr. Henry Chaney, Secretary Santa Barbara Museum of Natural History 2559 Puesta de Sol Road Santa Barbara, CA 93105—2936 We gratefully acknowledge the following individuals for their contributions to the Stohler Memorial Fund: Dr. Hans Bertsch, Dr. Rudiger Bieler, Dr. Henry W. Cha- ney, Mrs. Helen Dushane, Dr. Craig Edwards, Dr. William K. Emerson, Mrs. George P. Gerrodette, Dr. Terry M. Gosliner, Dr. Cadet Hand, Mrs. Carole M. Hertz, Dr. Car- ole S. Hickman, Dr. E G. Hochberg, Dr. John J. Holle- man, Dr. Richard I. Johnson, Ms. Kirstie Kaiser, Michael and Linda Kellogg, Dr. Artie L. Metcalf, Dr. Ellen J. Moore, Mrs. Marjorie Neiswanger, Dr. Larry C. Oglesby, Mr. William Pitt, Dr. Robert Robertson, Dr. Peter U. Rod- da, Dr. Barry Roth, Mrs. Carol Skoglund, Dr. Joan E. Steinberg, Dr. Walter A. Sunderland, Dr. John W. Tunnell Jr., Dr. Janice Voltzow, Dr. Shi-Kuei Wu. International Commission on Zoological Nomenclature The following Applications concerning mollusks were published on 31 March 2000 in volume 57, Part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on any of these applications is invited for publication in the Bulletin and should be sent to the Executive Secre- tary, LC.Z.N., c/o The Natural History Museum, Crom- well Road, London SW7 SBD, U.K. (e-mail: iczn@nhm.ac.uk). Case 2926. Trichia Lozek, 1956 (Mollusca): proposed emendation of spelling to Trichianinae, so removing the homonymy with Trichiidae Fleming, 1821 (Insecta, Coleoptera). Case 3086. Hyalinia villae adamii Westerlund, 1886 (cur- rently Oxychilus adamii,; Mollusca, Gastropoda): pro- posed conservation of the specific name adamii by re- placing the syntypes with a neotype. The following Applications concerning mollusks were published on 30 June 2000 in Volume 57, Part 2 of the Bulletin of Zoological Nomenclature. Case 3088. Doris verrucosa Linnaeus, 1758 (Mollusca, Gastropoda): proposed conservation of the generic and specific names by designation of a neotype. Case 3133. Peristernia Morch, 1852 (Mollusca, Gastro- poda): proposed conservation of Turbinella nassatula Lamarck, 1822 as the type species. The following Opinion concerning mollusks was pub- lished on 31 March 2000 in Volume 57, Part 1 of the Bulletin of Zoological Nomenclature. Copies of the fol- lowing Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., c/o The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e- mail: iczn@nhm.ac.uk). Opinion 1942. Haminoea [Turton] in Turton & Kingston in Carrington, 1830 and haminoeinae Pilsbry, 1895 (Mollusca, Gastropoda): placed on Official Lists as correct original spellings. The following Opinion concerning mollusks was pub- lished on 30 June 2000 in Volume 57, Part 2 of the Bul- letin of Zoological Nomenclature. Opinion 1950. Haliotis clathrata Reeve, 1846 (non Lich- tenstein, 1794) and H. elegans Philippi, 1844 (Mollus- ca, Gastropoda): specific names conserved. The Veliger 43(4):377—378 (October 2, 2000) THE VELIGER © CMS, Inc., 2000 BOOKS, PERIODICALS & PAMPHLETS Fossil Shells from Western Oregon—a Guide to Identification by ELLEN J. Moore. 2000. Chintimini Press, 3324 SW Chintimini Avenue, Corvallis, Oregon 97333, USA (http:// www.cmug.com/~chintimp). 131 pp., paperback, numer- ous illustrations. ISBN 0-9640066-1-8 US $12.00 This guidebook to Cenozoic fossil mollusks of western Oregon is the successor to two earlier, out-of-print guides by the same author. More than a simple guide to fossil shells, the first 43 pages include a brief history of pale- ontology in Oregon, an overview of geology in the state, how geology has formed the landscape, and instructions on how to recognize, collect, and identify fossils. Nine- teen more pages are devoted to self-guided geological field excursions in western Oregon. The introductory ge- ology sections and field trip guides are well illustrated. The core of the book begins with a useful guide to the morphology and terminology of mollusk shells. Fossil shells are then grouped by age from Eocene to Pleisto- cene, and are illustrated by photographs that accompany the species write-ups on each page. This is a useful device that avoids the pitfall of all too many guidebooks, where the photos are grouped onto plates that need to be book- marked with a thumb while you read the text. The quality of the fossil photos is very high, which mostly eliminates the need for morphological descriptions that are often the bane of guidebooks. Instead, the text accompanying each species’ photo gives sidelights into the usefulness of the species as a guide fossil for age, glimpses into its natural history, or comments on how it was preserved. An ex- tensive list of more technical literature for interested read- ers is included, as is a very complete index. In all, the thoroughness and clear presentation that make this guide- book a stand-out reflect the author’s long residence in the front ranks of West Coast research paleontologists. This an excellent book for the interested amateur, be- ginning student, or budding paleontologist who has ad- vanced beyond the Barney stage. Louie Marincovich, Jr. A Conchological Iconography G. T. Poppe & K. Grou, eds. Conchbooks, Hackenheim, Germany. Family Strombidae (1999): 60 pp., 130 pls. Text by K. KrerpL & G. T. Poppe. Plates by G. T. Poppe, L. MAN IN’T VELD & K. DE TurRcK. US$ 77.00. ISBN 3- 925919-29-5. Family Harpidae (1999): 69 pp., including 51 pls. G. T. Poppe, T. BRULET & S. P. DANcE. US$ 44.00. ISBN 3- 925919-28-7. Binder: US$ 11.00. ISBN 3-925919-27-9. The present work with the currently two published sets aims to provide a “high quality pictorial coverage of groups significant to collectors and scientific students alike.” It is a European production edited by Guido Pop- pe, who has written a number of conchological books (Poppe & Goto, 1991, 1992, 1993a, b), and Klaus Groh of the German publishing house ConchBooks (formerly Christa Hemmen). The format chosen is that of a loose leaf publication in A4 format held in a four-ring binder; although foreign to most Americans, both are standard in Europe. In comparison to Indo-Pacific Mollusca, the sets are printed throughout on heavier and glossy paper, which should withstand extensive usage. Each of the two fam- ilies contains a text part with unnumbered black and white text figures, and a set of plates with color images on black and white background. The individual images are on average 7 cm in size. The captions give locality data by country, size, and collection. Most specimens il- lustrated are in private collections. Additional plates show freak specimens, apices (Harpidae only), single dorsal and ventral 3 cm images of each species to provide an overview of the family’s diversity, and some live animals. The text of the Harpidae, including the genus Morum, is restricted to four pages of introductory remarks on the family, four pages of comparative notes on the 20 species discussed, including a partial synonymy list, and three pages of key references. A new species (Morum fatimae Poppe & Brulet in Poppe, Brulet & Dance, 1999) is also described. The first 41 plates illustrate the species. On average, approximately five specimens with eight images of each species are illustrated (Harpa amouretta with 14 specimens in 27 images). These are followed by four plates with apices, three overview plates, and three with live animals. The Strombidae set contains 11 pages of introductory remarks on the family. The individual species are treated at a pace of two to three (sub) species per page. The narrative contains information on range, size, and a de- scription of the shell, occasionally with some remarks in- cluding selected synonyms. Supraspecific taxa are not di- agnosed. An average of five specimens (11 for Terebel- lum terebellum Roding, 1798) are illustrated on 119 plates. Two additional plates contain juveniles and freaks, three for the overview, and six with live animals. Considering the main goal of the work to be a pictorial guide, the aim has certainly been achieved. Many more specimens are shown than in Walls’ (1980) Tibias, Page 378 Conchs, and Harps, and those illustrated were carefully chosen to illustrate as much intraspecific variability as possible. The quality of the plates is respectable, but not quite reaching the level of Anseeuw & Goto’s (1996) Pleurotomariidae book. Some plates of the sets I have available are somewhat dark. Critics may argue that too much printing space is taken up by the generous black background surrounding the shell specimens and the 1ii- lustrations of live animals. The selected bibliography. is useful for taxa recently described in some of the lesser known conchological publications, but does not provide many citations for older taxa. The matching of in-text citation to the bibliography is at times difficult due to the lack of adherence to any established citation format. Can Walls (1980) be relegated to the archive? I do not think so. For one thing, the narrative distribution infor- mation of the new series is not as detailed as Walls’ maps. The introductory remarks on the families are more com- prehensive in the work by Walls, who also provided di- agnoses for supraspecific taxa and a comprehensive syn- onoymy. These authors missed the opportunity to include this readily available information, or to provide further monographic data such as type locality, type specimen location, and type specimen status. The omission of di- agnoses of higher taxa is puzzling considering the 18 sub- genera recognized in the Strombidae, and the partisan dis- cussion of Mirabilistrombus Kronenberg, 1999, on page SD: The authorships of the individual sections are awk- The Veliger, Vol. 43, No. 4 ward. At the bottom of each page the authors of that particular page are indicated. Each of the sets has three fractioned collectives of authors for subsections of the text and the plates. Hopefully the editors will have the foresight to facilitate citation of subsequent sets by future authors. Overall, the work provides detailed image information on all species and many recently described subspecies within these two families. The casual collector may still do well with Walls (1980), but specialized collectors and institutional libraries may want to consider the purchase of these sets, particularly due to the first comprehensive treatment of the genus Morum. Further sets currently in preparation include Calliostoma (Trochidae), Ficidae, Haliotidae, Acavidae, Pectinidae I & II, and Turbinidae I & Il. Daniel L. Geiger Literature Cited ANSEEuUW, P. & Y. Goto. 1996. The Living Pleurotomariidae. Elle Scientific: Osaka. 202 pp. Poppe, G. T. & Y. Goto. 1991. European Seashells. Volume I. Christa Hemmen: Wiesbaden. 352 pp. Poppe, G. T. & Y. Goto. 1992. Volutes. Monstra Mondiale Ma- lacologia: Cupra Maritima. 348 pp. Poppe, G. T. & Y. Goto. 1993a. European Seashells. Volume II. Christa Hemmen: Wiesbaden. 221 pp. Poppe, G. T. & Y. GoTo. 1993b. Recent Angariidae. Monstra Mondiale Malacologia: Cupra Maritima. 32 pp., 10 pls. WALLS, J. G. 1980. Conchs, Tibias, and Harps. TFH Publications: Neptune, New Jersey. 191 pp. ELE Pe PLIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler (1901-2000), Founding Editor Volume 43 January 3, 2000 to October 2, 2000 TABLE OF CONTENTS January 3, 2000 The anatomy and systematics of Latiromitra, a genus of tropical deep-water Ptychatractinae (Gastropoda: Turbinellidae) PHILIPPE BOUCHET AND YURI I. KANTOR .............-. 1 Sex change, reproduction, and development of Crepidula adunca and Crepidula lingulata (Gastropoda: Calyptraeidae) RACHETs i GOLLING ceasceeeh oc eal oes eS Cue cke eee ome 24 Shell growth of Mytilus trossulus Gould, 1850, in Port Valdez, Alaska ARNY BLANCHARD AND HOWARD M. FEDER..........- 34 Two new Neogene species and the evolution of labral teeth in Concholepas Lamarck, 1801 (Neogastropoda: Muricoidea) THOMAS JE2DEMRIES@ pth ioe enone en med aro keine 43 Pisidium taraguyense and Pisidium pipoense, new species from northeastern Argentina (Bivalvia: Sphaeriidae) (ERISTIANSEMITUARTE eee a cea enere en Cney maureen sents cahcee ts 51 Development and metamorphosis of the planktotrophic larvae of the moon snail, Polinices lewisii (Gould, 1847) (Caenogas- tropoda: Naticoidea) ROBERTA V. K. PEDERSEN AND LOUISE R. PAGE ....... 58 Polygyrid land snails, Vespericola (Gastropoda: Pulmonata). 3. Three new species from northern California BARRY ROTH AND WALTER B. MILLER ............. 64 Distribution of the bonnet limpet, Hipponix conicus (Gastropoda: Hipponicidae), among host species in western Kyushu, Ja- pan KAZUNORI YAMAHIRA AND FUMITO YANO ..........-. 72 Growth and fecundity of Lymnaea elodes (Gastropoda: Lymnae- idae) under laboratory conditions LAUREN FLORIN, BERNARD FRIED, AND ADITYA REDDY .. 78 A new subspecies of the schoolmaster gonate squid, Berryteuthis magister (Cephalopoda: Gonatidae), from the Japan Sea OLEG NS KATUGIN] e356 ee ee eee 82 Arboreal Neritidae ROBERT H. COWIE AND BARRY D. SMITH ........... 98 Paleogerographic implications of late Paleocene Onestia onestae (Bivalvia: Cardiidae) in arctic Alaska JAY A. SCHNEIDER AND LOUIE N. MARINCOVICH, JR. .... 99 April 3, 2000 Nocturnalism in Aplysia oculifera (Adams & Reeve, 1850): an avoidance behavior minimizing exposure to ultraviolet ra- diation? TPATHPICAU Te sree eet eae cst crores te ole eirae cite aeie Meee nes 105 Spiomenia spiculata, gen. et sp. nov. (Aplacophora: Neomenio- morpha) collected from the deep waters of the West Euro- pean Basin PAMELA VARNOESK Ye aor cisneieeiiehens ese Gl cl aotsireieicstons 110 The buccinid gastropod Deussenia from Upper Cretaceous strata of California RICHARD L. SQUIRES AND LOUELLA R. SAUL ........ 118 Small, high-spired pulmonates from Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with description of a new genus, and with conservation statuses of 15 streptaxid species KENNETH C. EMBERTON AND TIMOTHY A. PEARCE .... 126 Three new Pacific species of Halgerda (Opisthobranchia: Nudi- branchia: Doridoidea) Cae CARESONFANDEPL US ELORR een iene ctom een iomenen 154 Taxonomic revision of the common Indo-West Pacific nudi- branch Phyllidia varicosa Lamarck, 1801 ALEXANDER FAHRNER AND MICHAEL SCHRODL ....... 164 Redescription and range extension of Bathydoris aioca Marcus & Marcus, 1962 (Nudibranchia: Gnathodoridoidea) ANGEL VALDES AND HANS BERTSCH .............. 172 Two new species of the family Hydrobiidae (Mollusca: Caeno- gastropoda) from Austria MARTIN HAASE, ERICH WEIGAND, AND HARALD HASEKE ee er eer N Rr ere gr id hc o6,.0 0 0,0.0 6 179 Additional data on the phanerobranch dorid Tambja simplex Or- tea & Motor, 1998 (Gastropoda: Nudribranchia: Polycera- tidae) JUAN LuCAS CERVERA, JOSE CARLOS GARCIA-GOMEZ, AND RICCARDO CATTANEO-VIETTI ........---- 190 The egg capsule and young of the gastropod Pyrulofustus dexius (Dall) (Buccinidae: Volutopsiinae) VLADIMIR: Vi \GULBIN) 20505 a eat eee ee 195 July 3, 2000 Revision of dorid Nudibranchia collected during the French Cape Horn Expedition in 1882—1883, with discussion of the ge- nus Geitodoris Bergh, 1891 MICGHABIE SS CHRODIE, 3) eras soba ooo a eter eee 197 Sex change in the hat snail, Calyptraea morbida (Reeve) (Gas- tropoda: Calyptraeidae): an analysis of substratum, size, and reproductive characteristics MING-Hul CHEN AND KERYEA SOONG ............. 210 Helicarionid snails of Mounts Mahermana, I[lapiry, and Vasiha, southeastern Madagascar KENNETH C. EMBERTON AND TIMOTHY A. PEARCE .... 218 Charopid snails of Mounts Mahermana, Ilapiry, and Vasiha, southeastern Madagascar, with description of a new genus and with conversation statuses of nine species KENNETH C. EMBERTON AND TIMOTHY A. PEARCE .... 248 Exploration of morphospace using procrustes analysis in stato- liths of cuttlefish and squid (Cephalopoda: Decabrachia) evolutionary aspects of form disparity JEAN-LOUIS DOMMERQUES, PASCAL NEIGE, AND SIGURD V. BOLETZIGY oie) ieo't aus, tec wi element eee eee eae ee 265 Shell size variation and aggregation behavior of Littoraria flava (Gastropoda: Littorinidae) on a southeastern Brazilian shore PAULO R. S. MOUTINHO AND CECILIA P. ALVES-CosTa 277 New information on a poorly known late Paleocene turrid gas- tropod from southern California and vicinity RICHARD); SQUIRES) 91-4 -aouc cee el ee nc eee 282 October 2, 2000 Dr. Rudolf Stohler (1901—2000) Memorial Issue ......... i Dr. Rudolf Stohler: some personal remembrances IRIANS IBERIIOES 5 ee apo os SOs Gen tale ee odors deo. oid ll Distribution, reproduction, and shell growth of limpets in Port Valdez, Alaska ARNY BLANCHARD AND HOWARD M. FEDER......... 289 Development and anatomy of Nitidiscala tincta (Carpenter, 1865) (Gastropoda: Epitoniidae) FRM CS OUGIN GH Sie hth Sein: ees Aw al beeen eke ae 302 The ecology and rapid spread of the terrestrial slug Boettgerilla pallens in Europe with reference to its recent discovery in North America HEIKE REISE, JOHN M. C. HUTCHINSON, ROBERT G. FORSYTH, AND TAMMERA J. FORSYTH .......... 313 Successful and unsuccessful predation of the gastropod Nucella lapillus (Muricidae) on the mussel Mytilus edulis from Maine GREGORWEES DIET Me rie ten enero isis st ct suey tee seven 319 Predation on juvenile Aplysia parvula and other small anaspi- dean, ascoglossan, and nudribranch gastropods by pycno- gonids C. N. RoGers, R. DE Nys, AND P. D. STEINBERG ..... 330 Life history of a hydroid/nudibranch association: a discrete-event simulation CHARLES M. CHESTER, ROY TURNER, MICHAEL CARLE, ANDEISARR Ye Gi FIARRIS| ee ia uence iene ene cna 338 Predation on the apple snail, Pomacea canaliculata (Ampullari- idae), by the Norway rat, Rattus norvegicus, in the field YOICHI YUSA, NAOYUKI SUGIURA, AND KATSUYA MEHINOSE ty bases) eee we, so Wiehest uae eatin, | eRe Gi 349 Three new species of dorid nudibranchs from southern Califor- nia, USA and the Baja California Peninsula, Mexico SANDRA V. MILLEN AND HANS BERTSCH ........... 354 A new species of hydrobiid snal of the genus Pyrgulopsis from northwestern Nevada ROBERT HERSHLER AND DONALD W. SADA .........- 367 AUTHOR INDEX IATEVES-GOSTAWG@URS as noe oes tin Cina aoe ROR 277 PENIN (BIEL)