iii et Ree keira i raipen wee UL ISSN 0042-3211 (THE VELIGER A Quarterly published by { 99 onne CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC MAR 69 cuve Berkeley, California “LIBR! R. Stohler (1901-2000), Founding Editor — Volume 50 March 11, 2008 Number 1 CONTENTS Anatomical Review and Preliminary Phylogeny of the Facelinid Nudibranchs (Opisthobran- chia: Aeolidina) of the Taxon Phyllodesmium Ehrenberg, 1831 DettsseE M. Ortiz AND TERRENCE M. GOSLINER Earliest Record of the Genus Haliotis (Mollusca: Gastropoda) from the Late Cretaceous (Cam- panian) of Los Angeles County, California GIN SEVeIe| GROVES AND?) OHING ME -ATEDERSON (ss mci aces ticles sie aoe cis elses sleveretie easter 24 Predatory Behavior and Diet of Eupleura sulcidentata Dall, 1890 (Gastropoda: Muricidae) from West Florida GRECORSH LIERBERTVAND) SHUBHABRATAUDAWIE. & oe an cw chelsea elec soha eve oe ela oe 27 A | arge New Species of Lobatus (Gastropoda: Strombidae) from the Neogene of the Domini- can Republic, with Notes on the Genus BERNARD M. Lanpau, Gis C. KRONENBERG, AND GREGORY S. HERBERT.............. 31 First Record of the Northeastern Pacific Patellogastropod Genus Acmaea from the Miocene of Japan and Its Paleobiogeographic Implications MOK OM MU RMTARAVAND MOMORTINASE onesie siete ole «isles onelepel ele nist eielcnnetele|clite iets ote.¢ 39 A New Phaenomenella Fraussen & Hadorn, 2006 (Gastropoda: Buccinidae), from the Anda- man Sea. INO ENG ERIAT SS EIN AT en petra ns Nir IN We aac is enter A ee EU ANA AT eat ce 48 Development of Tylodina fungina Gabb, 1865 (Gastropoda: Notaspidea) from the Pacific Coast of Panama IRUACTE BIE, CLO EIINT hig yer dic eh Amey lia te Secs Ae A Ain CH aIeaUesna ey ete a) nena SUR ee nee er 51 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. 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. THE VELIGER Scope of the Journal The Véeliger is an international, peer-reviewed scientific quarterly published by the California Malaco- zoological Society, a non-profit educational organization. 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Kellogg, City and County of San Francisco Christopher L. Kitting, California State University, Hayward David R. Lindberg, University of California, Berkeley Peter Roopharine, California Academy of Sciences Barry Roth, San Francisco Angel Valdés, Natural History Museum of Los Angeles County Geerat J. Vermeij, University of California, Davis Membership and Subscription Affiliate membership in the California Malacozoological Society is open to persons (not institutions) interested in any aspect of malacology. New members join the society by subscribing to The Veliger. Rates for Volume 50 are US $65.00 for affiliate members in North America (USA, Canada, and Mexico) and US $120.00 for libraries and other institutions. Rates to members outside of North America are US $75.00 and US $130.00 for libraries and other institutions. All rates include postage, by air to addresses outside of North America. Memberships and subscriptions are by Volume only. 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THE VELIGER 5 The Veliger 50(1):1-23 (March 11, 2008) © CMS, Inc., 2007 Anatomical Review and Preliminary Phylogeny of the Facelinid Nudibranchs (Opisthobranchia: Aeolidina) of the Taxon Phyllodesmium Ehrenberg, 1831 DELISSE M. ORTIZ School of Earth and Environmental Science, Washington State University Vancouver, 14204 NE Salmon Creek Avenue, Vancouver, WA 98686-9600, USA (e-mail: ortiz@vancouver.wsu.edu) TERRENCE M. GOSLINER Department of Invertebrate Zoology and Geology, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103, USA (e-mail: tgosliner@calacademy.org) Abstract. The anatomy and morphology of species of Phyllodesmium are described for P. parangatum Ortiz and Gosliner, 2003, P. magnum Rudman, 1991, P. poindimiei (Risbec, 1928), P. hyalinum (Ehrenberg, 1831), P. crypticum Rudman, 1981, P. serratum (Baba, 1949), P. colemani Rudman, 1991, P. kabiranum Baba, 1991, P. macphersonae Burn, 1962, P. briareum (Bergh, 1896), P. longicirrum (Bergh, 1905), P. pecten Rudman, 1981, P. opalescens Rudman, 1991, P. horridum (Macnae, 1954), P. iriomotense Baba, 1991 and P. guamensis Avila et al. A phylogenetic analysis supports the monophyly of Phyllodesmium. Species possessing an unbranched digestive tract are most basal, while more derived taxa have a highly ramified digestive tract. More basal species form a grade with symplesiomorphies such as an unbranched digestive tract, jaw with many denticles, absence of zooxanthellae and elongate foot corners. The remaining species have elaborate digestive tracts and have undergone morphological and physiological changes allowing the storage of zooxanthellae in the cerata, for use as a secondary food source. It is evident from this study that morphological changes occurring within Phyllodesmium correlate closely with their increased association with symbiotic zooxanthellae. This first parsimony-based phylogenetic study of Phyllodesmium largely supports the scenario of morphological evolution first proposed by Rudman (1991). INTRODUCTION Within the Aeolidina, the most diverse taxon is the Facelinidiae with more than 130 described species. One highly specialized group, species of Phyllodesmium Ehrenberg, 1831, has received considerable attention owing to the trophic specialization and evolution of symbiosis of members of this taxon. The facelinid taxon Phyllodesmium includes nineteen described spe- cies, almost all of which are known to be specialized predators on alcyonarian octocorals restricted to the Indo-Pacific tropics and adjacent temperate regions (Rudman, 1981b; Rudman, 1991; Gosliner et al, 1996; Avila et al., 1998; Ortiz, 2001; Ortiz & Gosliner, 2003; Burghardt & Wagele, 2004; Burghardt & Gosliner, 2006). The fact that most of these species also contain symbiotic zooxanthellae and exhibit a range of variation in the elaboration of digestive gland struc- tures to accommodate this symbiosis, makes them ideal candidates for phylogenetic and comparative biological studies. It needs to be determined if Phyllodesmium represents a monophyletic group and, if this proves to be the case, this particular study can illuminate the nature of the evolution of symbiosis within this particular clade. Species currently placed in Phyllodesmium have been placed in several different genera including Phestilla Bergh, 1874, Cratena Bergh, 1864, Hervia Bergh in Morch, 1871, Aeolidia Cuvier, 1798, Myrrhine Bergh, 1905, Favorinus Gray, 1850, Godiva Macnae, 1954, Ennoia Bergh, 1896, Phyllodesmiopsis Risso-Domin- guez, 1964, and Babiella Risso-Dominguez, 1964, reflecting the confusion surrounding the systematics of this group. Rudman (1981, 1991) summarized much of this confusion and the historical review will not be repeated here. Since the late 1800s, there has been a substantial amount of biological interest and research regarding ecological interactions and symbiosis between the dinoflagellate zooxanthellae Symbiodinium Freu- denthal, zooxanthellae, and a number of different marine hosts. This literature deals mostly with scler- The Veliger, Vol. 50;3Nom Figure 1. Living animals of Phyllodesmium species. A. P. colemani, photo by T.M. Gosliner. B. P. kabiranum, photo by R.F. Bolland. C. P. magnum, photo by T.M.Gosliner. D. P. macphersonae, photo by R.F. Bolland. E. P. parangatuwmn, photo by T.M. Gosliner. F. P. serratum, photo by R.F. Bolland. G. P. briareum, photo by T.M. Gosliner. H. P. crypticum, photo by T.M. Gosliner. I. P. poindimiei, photo by T.M. Gosliner. actinean corals and species of venerid clams in the taxon Tridacna Bruguiere (Fankboner, 1971; Goreau et al., 1973; Fitt and Trench, 1981; and many others), but has since expanded into the realm of nudibranch and coelenterate symbiosis. Two examples are Rud- man (1982), who worked on the aeolidoidean and arminoidean nudibranch mollusks and Kempf (1984) who studied species of Melibe Rang, Pteraeolidia Bergh and Berghia Trinchese. Hoegh-Guldbergh & Hinde (1986) also examined nudibranch-zooxanthellae symbiosis; they studied the effects of the presence of zooxanthellae on their nudibranch host. Recent works by Burghardt & Wagele (2004, 2006), Burghardt er al. (2005), and Burghardt & Gosliner (2006) have examined photosynthetic activity in a variety of additional opisthobranchs, including several species of Phyllodesmium. The available literature, however, has dealt mostly with associations regarding nudibranch and dinofla- gellate symbiosis in a descriptive form. Research on the evolution of aeolid nudibranchs and their specific hosts is virtually absent from the literature. Aside from the work of Rudman (198la, 1981b, 1982, 1987, 1991), publications regarding evolutionary adaptations as a result of morphological modifica- tions to accommodate the respective symbionts are rare. No phlylogenetic study based on the ecological interaction between a facelinid nudibranch taxon and its host has been published. The preliminary phylogeny of the facelinid nudibranchs belonging to the taxon Phyllodesmium Ehrenberg, 1831 is the first such study. The various adaptations and other anatomical variations, which have evolved in species of Phyllo- desmium, have resulted in it becoming one of the most morphologically diverse in the Aeolidina (Figure 1; Rudman, 1991). The reconstruction of phylogenies within the aeolid nudibranchs has been problematic. Difficulties have been encountered by those systema- tists that have attempted to clarify the phylogeny of aeolids (Miller, 1974; Gosliner and Ghiselin, 1984). Historically, the branching of the digestive gland, reflected in the ceratal arrangement, and the position of the anus, has been greatly emphasized in the D. M. Ortiz & T. M. Gosliner, 2007 classification of the Aeolidina (Odhner, 1934). Miller (1971) also considered the “‘branching of the digestive gland and position of the anus to be main features for delineating genera.” The taxon Phyllodesmium, as well as its nominal species, have been delineated and described based on these characters. However, Rud- man (1991) related ceratal arrangement, digestive gland branching, and other major interspecific mor- phological characters to the ecology of their food sources. The purpose of this study is to fully review and supplement the anatomy of species of Phyllodesmium in order to produce a preliminary phylogenetic analysis. The results of this study can be used as a basis for examining morphological changes within Phy/llodes- mium in response to coevolution with zooxanthellae symbiosis. MATERIAL AND METHODS Morphological Studies Specimens of previously described species of Phy/lo- desmium, accompanied by color slides of the living animals, were the primary source for morphological characters used in this study. Most importantly, examination of the material housed at the California Academy of Sciences (CASIZ) and the Australian Museum (AM) provided a wealth of specimens that were used to assess and verify doubtful and uncertain characters, as well as developing new characters for the phylogenetic analysis. Species of Facelinidae were examined and compared morphologically, using reproductive and radular mor- phology, and anatomical features including ceratal arrangement and structure, location of anus, branching of ceratal digestive gland, and shape of the anterolat- eral foot corners. Individual specimens were dissected to examine detailed structure of the cerata, buccal mass, and reproductive system. Dissections and scale drawings were made using a dissecting microscope with a camera lucida. An incision was made along the entire midline of the foot. The reproductive systems, as well as external features (e.g., rhinophores, anterior and posterior foot, location of anus, ceratal arrangement, etc.) were then examined. A LEO series 1400 Scanning Electron Microscope (SEM) at the California Academy of Sciences (CAS) was used to make scaled digital pictures of the structure of the radulae and jaws, in order to survey the phylogenetic characters used in the study. The cerata were dissected and stained. An average of 3 to 4 cerata were extracted from the dorsum of some of the specimens representing several species. Micro- graphs of the ceratal digestive branching were digitally captured using a Kodak MDS 100 camera Page 3 mounted on an M400 Wild microscope. Some of the more problematic cerata were drawn to scale (using a camera lucida on a Nikon SMC-10 dissecting micro- scope) or photographed (using an FX-35 DX Nikon camera mounted on an SMZ-U Nikon dissecting microscope). The cerata of one species, Phyllodesmium crypticum, were stained with a solution of acid fuchsin and 70% ethanol, dehydrated in alcohol, cleared with xylene, and mounted in Permount on a microscope slide. Phylogenetic Analysis Taxa. For the phylogenetic analysis, 20 taxa have been considered (Table 1). In order to determine polarity of morphological change within Phyllodes- mium, specimens of Godiva quadricolor (Barnard, 1927) and Favorinus japonicus Baba, 1949, were selected as outgroup taxa based on the fact that they represent relatively underived members of the Facelinidae (Willan, 1987). In the absence of a more comprehensive phylogeny of the Facelinidae, we agree with Willan (1987) that these taxa represent appropriate outgroups for polarizing characters within Phyllodesmium. These data were compared with the descriptions of G. quadricolor from Willan (1987) and F. japonicus from Rudman (1980). Phylogenetic Methods To develop a phylogenetic hypothesis for Phyllodes- mium, the morphological data were entered into a data matrix using MacClade 3.01 (Maddison and Maddi- son, 1992). All the characters used were assigned equal weight and treated as unordered. PAUP 4.0b4a (Swofford, 2000) was used for phylogenetic reconstruc- tion using a heuristic search with the TBR branch swapping option. One hundred random start trees were obtained by stepwise addition. Three characters were deleted in the subsequent analyses due to being uninformative and ambiguous (14, 21 and 30). The deleted characters are indicated in parenthesis in the character description section. Bremer decay analyses were performed by subsequent analysis with a series of iterations that examined successive trees, each one step longer, to estimate branch support using the method- ology of Bremer (1994). Characters. The 31 characters used to resolve the phylogeny of Phy/lodesmium are listed in Table 2. Of these characters, twenty-nine are binary and two are multistate. The characters states are represented with numbers: 0, ““~presumed”’ plesiomorphic condition; 1-2, “presumed” apomorphic conditions (refer to Table 3). Page 4 The Veliger, Vol-50; Nom Table 1 Sources used to describe the species in the present study. Abbreviations: R80, (Rudman, 1980); R81b, (Rudman, 1981b); R91, (Rudman, 1991); B37, (Baba, 1937); B91la,b, (Baba, 1991a,b); Bu6é2, (Burn, 1962); M54, (Macnae, 1954); Rb28, (Risbec, 1928); BeOS, (Bergh, 1905); C98, (C. Avila et al., 1998); RC87, (R.C. Willan, 1987); E31 (Ehrenberg, 1831); Br04 (Burghardt et al., 2004); Br06 (Burghardt 2006); T.G., Terrence Gosliner; R.F.B., Robert F. Bolland; PNG, Papua New Guinea; |.r., literature review; p.p., published pictures; p.s., present study; N/A, Phyllodesmium species CASIZ Accesion # Slide # Reference Type of research P. poindimiei 086009, 93947 T.G.: 086009 Rb28, R91, p.s. Re-examined P. parangatum T.G.: 106472,105657, 96325, T.G.: 105676, p.s. New Species 103702, 105676 106472,96325, 105657 P. magnum 79239,79221 PNG-1988 R91, p.s. Re-examined P. hyalinum 69970, 68731 R.F.B 2161 E31, BeOS, B37, R81b, R91, Re-examined MS4, p.s. P. horridum N/A p-p. R81b, R91, M54 Lr. P. crypticum 99314, 106465 Station 35 Ligpo R81b, R91, p.s. Re-examined P. serratum 114759 P. serratum/Okinawa B9la, R91, p.s. Re-examined P. colemani 110358 T.G.: 110358 R91, p.s. Re-examined P. kabiranum 89035, 103793 R.F.B. 3158 B9I1b, p.s. Re-examined P. macphersonae 115724, 104700, 65346 R.F.B. 3304 R91, R8&1b, Bué2, p.s. Re-examined P. longicirrum N/A p.p. R81b, R91 Lr. P. pecten N/A p-p. R81b IL ir P. opalescens N/A p.p. R91 Ieee P. iriometense N/A p.p. B91b Lr. P. guamensis N/A p-p. C98 ie, P. briareum 65346, 65299, 83678 PNG-1988 R91, p.s. Re-examined P. jakobsenae N/A p-p Br04 Lr P. rudmani N/A p-p Br06 Lr G. quadricolor N/A p-p. RC87, B27 Ive F. japonicus N/A p-p R80, B49 Lr. Non Applicable. SYSTEMATIC DESCRIPTIONS Introductory Remarks Species descriptions include anatomical information derived from the present study and from literature review. The anatomical data derived from _ other references have been repeated for a comparison with the data obtained from this study. The data for the re- examined specimens is from original research for the present study, unless otherwise stated in the text. Some morphological characters, such as the distribution and storage of zooxanthellae, could not be determined by the dissections performed, hence literature review was necessary. Family Facelinidae Bergh, 1889 Phyllodesmium Ehrenberg, 1831 Phyllodesmium Ehrenberg, 1831 [type species by subsequent designation (Gray, 1847), Phyllodesmium i/inum Ehrenberg, 1831] hi Sergh, 1905 [type species by monotypy, gicirra Bergh, 1905] Bai usso-Dominguez, 1964 [type species by otypy, : serrata Baba, 1964] Phyllodesmiopsis Risso Dominguez, 1964 [type species by monotypy, Favorinus horridus Macnae, 1954] Diagnosis: Alcyonarian-eating aeolids with cerata capable of being autotomized. Cerata slightly or extremely flattened, lacking functional cnidosacs (a synapomorphy for Phy/lodesmium). Oral glands absent with a pair of discrete tubular salivary glands present. Rhinophores smooth or slightly nodular (a synapo- morphy for Phyllodesmium). Foot corners angular (a synapomorphy for Phyllodesmium). Ceratal arrange- ment variable. Pre-cardiac cerata arranged in single or double rows, while the post-cardiac cerata arranged in single or double rowed arches, simple rows, or a mixture of simple row arches and simple rows. Cleioproctic anus present in first post-cardiac arch or behind first postcardiac arch. Masticatory border or jaw with a single row of denticles or smooth (a synapomorphy for Phyllodesmium). Radular formula 0.1.0. Teeth usually having long pointed central cusp or reduced one. Each cusp with lateral flange along each side, with or without denticles. Genital opening below anterior limb of first right pre-cardiac arch or row. Reproductive system with single receptaculum seminis. D. M. Ortiz & T. M. Gosliner, 2007 Page 5 Table 2 Character descriptions and character states of present study. 1. Body size 12. Anterior foot corners 0 = moderate 0 = elongate 1 = large 1 = angular 2. Branching of digestive gland postcardiac group 23. Radular denticle location 0 = along masticatory border of tooth 1 = extent to central cusp of tooth 13. Vertical position of anus on first 24. Base of teeth 0 = absence of branched duct 0 = posterior 0 = long 1 = slightly branched duct 1 = dorsally 1 = short 2 = highly branched duct 3. Storage of zooxanthellae 14. Rhinophore size 25. Radular base of teeth 0 =absent 0 = long 0 = long 1 = present 1 = extended 1 = short 4. Ceratal surface 15. Rhinophore surface 26. Denticle size on jaw plates 0 = smooth 0 = swelling on lamaelle 0 = small 1 = nodular 1 = smooth or slightly nodular 1 = large 5. Ceratal shape 16. Masticatory border of jaw 27. Cnidosac 0 = cylindrical 1 = flatenned 2 = smooth 0 = several rows of denticles 1 = single row of denticles 0 = present 1 = absent 6. Ceratal apex 17. Number of denticles 28. Penial spine 0 = blunt 0 = numerous 0 = present 1 = curled 1 = few and elongate 1 = absent 7. Ceratal arch in precardiac group 18. Cusp of teeth 29. Female gland mass 0 = present 0 = short 0 = elongate 1 = absent 1 = long 1 = bulbous 8. First postcardiac arrangement 19. Radular denticles 30. Penial complex 0 = arches 0 = present 0 = large 1 = rows 1 = absent 1 = small 9. Second postcardiac arrangement 20. Radular denticle arrangement 31. Arrangement of radular denticles 0 = arches 0 = separated 0 = single rows 1 = single or double rows 1 = tightly congested 1 = double rows 10. Third postcardiac arrangement 21. Radular denticle length 0 = arches 0 = long 1 = single or double rows 1 = short 11. Foot width 22. Radular denticle tip 0 = wide 0 = blunt 1 = narrow 1 = pointed Prostate forming gland mass at the base of penis. Penis simple, unarmed. Phyllodesmium parangatum Ortiz & Gosliner, 2003 (Fig. 1E) Material examined: Holotype, California Academy of Sciences, CASIZ 106472 near Twin Rocks, Anilao, Batangas Province, Luzon, Philippines, 15 April 1996, T. Gosliner. Paratypes: three specs, one dissected, CASIZ 105657, Devil’s Point, Maricaban Island, Batangas Province, Luzon, Philippines, 25.3 m depth, 23 February 1995, T. Gosliner. One specimen dissected, CASIZ 105676, Bonito Island, Maricaban Island, Batangas Province, Luzon, Philippines, 2.1 m depth, 27 February 1995, T. Gosliner. One specimen, CASIZ 96325, Cemetary Beach, Maricaban Island, Batangas Province, Luzon, Philippines, 13 March 1994, T. Gosliner. Distribution: So far, known only from the original localities in the Philippines. Remarks: The anatomy of this species is completely described by Ortiz and Gosliner (2003). Phyllodesmium magnum Rudman, 1991 (Figs. 1C, 2A—C) Phyllodesmium sp. Orr, 1981: 69. ?Phyllodesmium sp. Willan and Coleman, 1984: 48, fig. 154. Phyllodesmium magnum Rudman, 1991: 190-193, figs. 20A, B, 22-24, 27C, D, 28, 29. Material examined: Two specimens, one dissected, CASIZ 79239, Horseshoe Cliffs, 1 km WNW of Onna Page 6 The Veliger, Vol. 50, No. 1 Table 3 Character states present in Phyllodesmium species. Data code: see Table 2. Abbreviation: ?, no data available. _ No Ww Nn OV . horridum . serratum . poindimiei opalescens . briareum colemani . magnum . hyalinum crypticum . macphersonae longicirrum pecten iriomotense . kabiranum . parangatum . guamensis . quadricolor japonicus . jakobseni . rudmani Rel ash eea PAP aio} ashawleneselerche ae) sh eas) eas each ash ele} mops) KB ROOFPrFRrF OOH OoOOCOFCOCCoCCO NNOCONNNKNNNNNNNN CHK OC KB ROOR Ve eR ee eee RR OOOO SCoOFPooOCoOOFCOFH OCC COCSO!]A SCKHOOrPKeReeHRHOoOHePHeHHoocoocS CFP CORR RP ORF EP Ee RH OOOH CSO KB VOOR ROD OCOCOCOCOFRPORCOOCSO]A SO OS SO Ol OS eS Oo OHS SS So = 'S S| 00 = i) = — — ~~) Ww = £ = n = a = ~ = foe) SO OS Ole St SO On Se Ore) | 6 hte eS Sey Ss Ss SS SS Sa SS) eR OF NR RKP RRP NNR KEP NNNK Ne BE OO DR eee aD RP EP DD VDOrFOCO SOOO COOKFCOoOHKPOOOKFHOHCSO KB ROORP FPR OP eee eRe EER OF Seooceeecoeoionoeoaooace SSOCDOFCCOCORP HP ORK HH OOOO Sao Qin SS SSS SSS SS aS SS ae KBR OORP ER OR RP eR eRe RR OR RE \o i) So NO — N N NO Ww tw £ tw n to ron tw XN tw lon) i) Ne) Ww So Oe) = . horridum . serratum . poindimiei opalescens briareum colemani . magnum . hyalinum crypticum . macphersonae longicirrum pecten irlomotense . kabiranum parangatum . guamensis . quadricolor . Japonicus . jakobseni . rudmani SOLUS TEG CULO UL AUAULaU Uh AO: SUSU aU CU eUmO! SUay ecooocoeooorFaooOHFcCcooHCS KH RKCOFPRrFCOH- HK COOFR COOPER BH OOHOoroorrreHeeHrRHoooo FBrFPOOrFROOOKRRFR ORF OR HF OOH BPrFPooreorocoorHe Hee HBHHroooso eRe RP OR RK Re RK Se KF OF OR Se KEK ORF ee eS FPrPoOoOwrr rere KH OCOCOF OCOOCOCrFK OO ee > eC CC CC nC ion ii Wan ia) mR ODDO Re RB Re SE EK OR See TS) Sey SS) Sree) CS) ye Oy) ©) SOW RFP FOr DP RP OR RP RP FOr ew L SCOnyrOr COR OF KP OF OO OrF ae AOS OLOlLOrOo OOO OOo OOS Village, Okinawa, Ryukyu Islands, Japan, 1.5 m depth, 6 May 1991, R.F. Bolland. One specimen, dissected, CASIZ 79221, Horseshoe Cliffs, 1 km WNW of Onna Village, Okinawa, Ryukyu Islands, Japan, 6 May 1991, R.F. Bolland. Distribution: Type locality is New Caledonia, however its distribution is widespread. Northern Western Australia, Marshall Islands and Hong Kong (Rudman, 1991). Collected from Tanzania, Papua New Guinea, yimes and Japan (Gosliner et al., 1996; present External morphology (Fig. 1C): Body large, broad, up to 120 mm (Rudman, 1991). Present specimens (CA- SIZ 79239 and CASIZ 79221) 45 mm and 68 mm in length, respectively; elongate body extending broadly from the anterior end, tapering at posterior end. Foot corners, long, angular. Oral tentacles, rhinophores slender, smooth. Cerata large, smooth, flattened and curved apically, extending all along the animal’s dorsum. Ceratal arrangement consisting of single vertical rows on distinctive ridges, double row of precardiac cerata along each side of body. Reproduc- tive opening below right-sided double precardiac rows. D. M. Ortiz & T. M. Gosliner, 2007 Page 7 Figure 2. Phyllodesmium magnum. A. Lateral view of radula. B. View of digestive tract in a ceras. C. Reproductive System. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 0.50 mm: B = 10 mm: C = 5 mm. Renal opening on right side and centred within interhepatic space. Anal papilla between right post- cardiac cerata and behind uppermost cerata of first posteardiac row on prominent mound. Up to Il postcardiac rows present with up to 8 or more cerata per row. Cerata and digestive gland (Fig. 2B): Cerata flattened, providing increase of surface area, allowing for the branching of the digestive gland duct on each ceras. Digestive gland extending all along cerata with numerous branches diverging into many secondary and tertiary branches. Flattened branches with terminal chambers; zooxanthellae present in parts of cerata exposed to sunlight, including digestive gland in both the body wall and foot (Rudman, 1991). Buccal armature (Fig. 2A): Radular formula of 45 mm long specimen 21 X 0.1.0 (CASIZ 79239). Tooth short, with wide base enclosing part of basal posterior structure. Cusp elongate, pointed. Denticles small, short, continuous, extending halfway down each tooth. Jaw with smooth masticatory border (Rudman, 1991). Reproductive system (Fig. 2C): Preampullary duct long, narrow, extending into a broad, long folded ampulla via thin duct. Duct bifurcating into recepta- culum seminis and penial complex. Penial complex extending into short, narrow prostate, connecting to membrane gland via small opening. Prostate with overlying layer of tissue, connecting with a massive, folded and bulbous female gland mass. Phyllodesmium poindimiei (Risbec, 1928) (Figs. 11, 3A—C) ?Phidiana tenuis Eliot, 1905: 287-288. Page 8 The Veliger, Vol. 50, No. 1 fmg Figure 3. Phyllodesmium poindimiei. A. View of digestive tract inside a ceras. B. Radular denticles. C. Reproductive system. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 10 mm; B = 0.002 mm; C = 5 mm. Aeolidia poindimiei Risbec, 1928: 246-247, fig. 78, Pl. 9, feast Phestilla poindimiei 96. Phyllodesmium poindimiei — Rudman, 1981b: 224-229, figs. 3-7, 21B, 25C, 27. Risbec, 1953: 138-139, figs. 93— Material examined: Three specimens, CASIZ 86009, Devil’s Point, NW side of Maricaban Island, Batangas, Luzon Island, Philippines, 25.3 m depth, 26 March 1993, T.M. Gosliner. One specimen, CASIZ 105746, Sepok, Maricaban Island, Batangas Province, Luzon, hilippines, 16.7 m depth, 24 February 1995, T.M. lin One specimen, dissected, CASIZ 93947, Watubela Islands, E Banda Sea, Moluccas, Indonesia, 12.1 m depth, 11 November 1993, P. Fiene. Distribution: Originally described from New Caledonia (Risbec, 1928). Found and redescribed from New South Wales and Western Australia (Rudman, 1991). Specimens also collected from Indonesia, Japan, and the Philippines (present study). External morphology (Fig. 11): Body of moderate size, elongate, 13.5mm in length. Body shape wider anteriorly, extending narrowly from anterior to poste- rior end. Anterior foot narrow anteriorly. Foot corners angular. Rhinophores moderately long, smooth, ap- proximately equal in length size to oral tentacles. Cerata cylindrical, smooth and curved, similar to P. D. M. Ortiz & T. M. Gosliner, 2007 magnum and P. parangatum. Dorsum partially exposed, not covered by numerous recurved cerata. Ceratal arrangement, except for the precardiac arch, of single rows with up to 3 to 5 recurved cerata each, on right side of body. Both renal and reproductive openings immediately below precardiac arch, in close proximity to first postcardiac row within interhepatic space. Anal papilla immediately below first postcardiac row. Postcardiac rows ranging from 7 to 9 rows on each side of dorsum. Cerata and digestive gland (Fig. 3A): Ceratal digestive gland duct extending to apical end of each ceras, branching outwardly in close proximity to ceratal wall. Simple branching ducts perpendicular to central duct, extending in terminal sac in variable manner. Cerata cylindrical, smooth, terminating in curled apex. Buccal armature (Fig. 3B): Radular formula 24 X 0.1.0 (CASIZ 93947). Tooth broad with long base, extending anteriorly and encasing posterior edge of adjacent tooth. Cusp long, with denticulation, extending along centre edge of margin. Degree of separation between elongate, adjacent denticles variable. Masticatory border of jaw smooth (Rudman, 1991). Reproductive system (Fig. 3C): Preampullary duct long, broad, extending into bulbous ampulla via thin duct. Duct bifurcating into receptaculum seminis and into penial complex. Penial complex extending into large, short, bulbous prostate. Connection between albumen and membrane glands basal. Prostate con- necting with folded. Female gland mass long, massive. Remarks: Rudman (1981) considered the likelihood that Phidiana tenuis Eliot, 1905) may be a senior synonym of Phyllodesmium poindimiei based on simi- larities of external morphology and radula. He was reluctant to place these taxa in formal synonymy, pending the collection of additional material from East Africa, the type locality of P. tenuis. We concur with this approach to resolving the nomenclature of this species. Phyllodesmium hyalinum Ehrenberg, 1831 (Fig. 4A—D) Phyllodesmium hyalinum Ehrenberg, 1831: 32. Phyllodesmium xeniae Gohar & Aboul-Ela, 1957: 131— 144, P1.1 Favorinus horridus brevitentaculatus Engel and van Eeken, 1962: 28-29, fig. 5. Phyllodesmium orientale Baba, 1991b:109, figs. 1-3 ; pl. 1, fig. 1, possible synonym. Material examined: One specimen, CASIZ 69970, Seragaki Beach, 1.3 km ENE of Maeki-zaki, Okinawa, Ryukyu Islands, Japan, 1.5m depth, 13 May 1989, Page 9 R.F. Bolland. Two specimens, one dissected, CASIZ 68731, Hole in the Wall, near Hussein Village, north of Madang, north coast, Papua New Guinea, 6.4 m depth, 21 July 1989, T.M. Gosliner. Distribution: First described from the Red Sea (Ehren- berg, 1831), but has been recorded more recently from the Philippines (Bergh, 1905), Japan (Baba, 1937), Tanzania (Rudman, 1981b), South Africa (Gosliner, 1987), Papua New Guinea, Japan, Tanzania, Indonesia (present study). External morphology: Body of moderate size, 10.3 mm in length. Foot wide, extending posteriorly, tapering into reduced posterior end. Oral tentacles long, about equal to the length of smooth rhinophores. Curved cerata numerous, covering surface of dorsum. Ceratal arrangement in arches in precardiac and postcardiac cerata. Number of cerata per arch 7 to 10 on each arch on each side of dorsum. Reproductive opening situated at base of precardiac arch, on right side of dorsum. Renal opening centred in the interhepatic space, in close proximity to first postcardiac arch. Anal papilla on a distinctive mound located on right side of the dorsum, outside arch on the posterior side of the first postcardiac arch. Up to six postcardiac ceratal arches present on each side of the body. Cerata and digestive gland (Fig. 4C): Digestive gland extending entire length of ceras with secondary and tertiary branches extending in ““web-like form” parallel to ceratal wall. Ceratal surface nodular. Branches terminating in small chambers, assumed to be the storage areas for the zooxanthellae from its alcyona- cean feeding source, Xenia spp. (Rudman, 1991). Nodular cerata flattened and curled apically. Buccal armature (Figs. 44A—B): Radular formula 25 < 0.1.0 (CASIZ 68731). Tooth narrow with long base, elongate cusp. Denticles visible halfway down tooth. Denticles short, pointed, well separated. Jaws with single row of large, sparse denticles all along mastica- tory border (Rudman, 1991). Reproductive system (Fig. 4D): Preampullary duct short, expanding into elongate, bulbous ampulla via narrow duct. Duct bifurcating into receptaculum seminis and penial complex. Penial complex extending into large, elongate, folded prostate. Prostate connect- ing to folded albumen gland, with surrounding membrane gland, and massive and long female gland mass. Remarks: Phyllodesmium orientale Baba, 1991 was described as a distinct species (Baba, 1991b) based on two specimens collected from Japan. Baba noted that it was similar to P. hyalinum, except that it has an anal position that is more similar to P. crypticum. However, Page 10 The Veliger, Vol. 50, No. 1 Figure 4. Phyllodesmium hyalinum. A. Denticles on masticatory border of radula. B. Anterior view of radular tooth. C. View of digestive tract inside a ceras. D. Reproductive system. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 0.010 mm; B = 0.003 mm; C = 5 mm; D = 5 mm. D. M. Ortiz & T. M. Gosliner, 2007 Page 11 Figure 5. Phyllodesmium crypticum. A. View of digestive tract inside a ceras. B. Lateral view of radula. C. Reproductive system. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 10 mm: B = 0.05 mm: C = 5 mm. it is evident from Baba’s illustration (pl. 1, fig. 1) that the anus is rather dorsal in its position, similar to P. hyalinum. We suspect that P. orientale is synonymous with P. hyalinum and list it as a probable synonym, pending discovery of more material from Japan. Phyllodesmium crypticum Rudman, 1981 (Figs. 1H, 5A—C) Phyllodesmium cryptica Rudman, 1981b: 232—236, 244— 261, figs. 10OA, B, 11, 12D, 13, 14B, 22B, 23, 26B, 27; Rudman, 1991: 194, fig. 28. Material examined: One specimen, dissected, CASIZ 99314, Huamja Island, NE side Manua, Mtwara Region, Tanzania, 4 November 1994, T.M. Gosliner. Two specimens, one dissected, CASIZ 106465, near Twin Rocks, Batangas Province, Luzon, Philippines, 9.1 m depth, 15 April 1996, T.M. Gosliner. Distribution: Originally described from Dar es Salaam, Tanzania (Rudman, 1981b). Recorded from New South Wales and Western Australia (Rudman, 1991). Specimens collected from Philippines and Japan (present study). External morphology (Fig. 1H): Body of moderate size, 10mm in length. Foot wide, elongate, tapered at posterior end. Anterior foot corners tapered. Rhino- phores, oral tentacles smooth and long; rhinophores shorter than oral tentacles. Cerata numerous often obscuring dorsum of body. Ceratal arrangement in arches. Number of cerata undetermined due to detachment of most of the cerata of the material examined. Up to 7 postcardiac arches present on each Page 12 ihe Veliger, Wolk 50; Nomi side of the body. Reproductive opening situated below anteriormost portion of right precardiac arch. Renal opening immediately in front of uppermost portion of the right-sided first postcardiac arch. Anal papilla located within first postcardiac arch, on right side of body. Cerata and digestive gland (Fig. 5A): Digestive gland extending all along ceras. Ceratal surface nodular with secondary and tertiary branches extending in a ““web- like form” to the ceratal wall. Branches terminating in small chambers, assumed to be the storage area for the zooxanthellae extracted from its alcyonacean prey, Xenia spp. (Rudman, 1991). Cerata flattened, curved at apex. Buccal armature (Fig. 5B): Radular formula 24 x 0.1.0 (CASIZ 106465). Tooth with broad, elongate base, extending anteriorly to adjacent tooth. Cusp elongate anteriorly. Short, pointed, well-separated denticles present on all teeth. Single row of large denticles present on masticatory border of jaw plates. Reproductive system (Fig. 5C): Preampullary duct short, extending into long, folded, bulbous ampulla. Ampulla extending via broad duct to receptaculum seminis and penial complex. Penial complex small, with proximal end connecting to massive and_ folded albumen and membrane glands. Distal end connecting with small, irregular and bulbous female gland mass. Remarks: Phyllodesmium hyalinum and P. crypticum, may easily be confused as the same species, but differ in the base of the teeth, anal position, rhinophore size and size of penial complex. Phyllodesmium hyalinum has greatly extended rhinophores, an anus located above the first postcardiac arch, a narrow base and a large penial complex. Phyllodesmium crypticum has moder- ately long rhinophores. The anus 1s inside the first post- cardiac arch. The radular teeth have a wide base and the penial complex is small. Phyllodesmium serratum (Baba, 1949) (Figs. 1F, 6A—C) Hervia serrata Baba, 1949: 105-106, 179, pl. 46, figs. 156-157, text figs. 142-143. Cratena serrata — Baba, 1955: 36, 56. Babiella serrata — Risso-Dominguez, 1964: 223. Phyllodesmium serrata — Rudman, 1981b: 260. Phyllodesmium serratum — Baba, 1991a: 101, figs. A— Eiiplelshigsl=3): Material examined: One specimen dissected, CASIZ 114759, | km WNW of Onna Village, Horseshoe Cliffs, Okina\ Ryukyu Islands, Japan, 44.2 m depth, 18 May 19 R.F. Bolland. Distribution: Originally described from Japan (Baba, 1949). Recorded from different regions of Australia: Victoria, New South Wales and parts of the Northern Territory (Rudman, 1991). External morphology (Fig. 1F): Body, 32 mm in length, moderate size. Foot wide and elongate, tapering posteriorly, tapered anterior foot corners. Rhinophores smooth and moderately long, as long as the oral tentacles. Cerata, smooth, long, straight, extending across whole body, arranged in arches, precardiac arch can have up to five cerata on each side, postcardiac arches range from 7 to 10 arches on each side, each arch containing up to 7 cerata on each arch. Reproductive opening between center and uppermost edge below precardiac arch. Renal opening in front of first postcardiac arch. Anal papilla on a distinctive mound, centred inside first postcardiac arch. Cerata and digestive gland (Fig. 6A): Branched diges- tive gland, extending as finger-like projection inside cerata, short lateral branches evident, cerata are cylindrical, smooth, long and numerous. Buccal armature (Figs. 6B—C): Radular formula 24 x 0.1.0 (CASIZ 114759). Tooth wide with short base, extending anteriorly and covering the posterior basal edge of the tooth in front, long cusp, denticulation extending all along border of each tooth. Tightly joined and long denticles, extending into pointed tip, small denticles on masticatory border of jaw plates (Baba, 1991a). Phyllodesmium colemani Rudman, 1991 (Figs. 1A, 7A—B) Phyllodesmium sp. Coleman, 1988: 14-15, 1989: 7, 54. Phyllodesmium colemani Rudman, 1991: 187-190, figs. 14C, 18, 19, 20C—-E, 21, 27-29. Material examined: Two specimens, one dissected, CASIZ 110358, Bus Stop Reef, Balayan Bay, Batangas Province, Luzon Island, Philippines, 23 April 1997, M. Miller. Distribution: Known from its type locality, Lord Howe Island, Coral Sea (Rudman, 1991) and the Philippines (present study). External morphology (Fig. 1A): Body 22.5 mm in length, of moderate size, extending uniformly narrowly from anterior to posterior end. Foot narrow and elongate. Rhinophores greatly extended and smooth, similar to oral tentacles. Cerata long, smooth, flat- tened, extending along dorsum of body, visible dorsum. Ceratal arrangement with single postcardiac vertical rows and precardiac arch, up to 3 to 4 cerata on each of the 7 to 8 postcardiac rows located on each side of body, up to 3 to 5 cerata on each side on precardiac D. M. Ortiz & T. M. Gosliner, 2007 A dt Page 13 Figure 6. Phyllodesmium serratum. A. View of digestive tract inside a ceras. B. Lateral view of radula and its denticles. C. Lateral profile of radula. Abbreviations: dt, digestive tract. Scale bars: A = 5 mm; B = 0.02 mm; C = 0.07 mm. arch. Reproductive opening, renal opening, and anal papilla located on right side of body. Reproductive opening located below anterior-most portion of pre- cardiac arch, renal opening located in center of interhepatic space; anal papilla raised on a distinctive mound, located outside first postcardiac row. Cerata and digestive gland (Fig. 7A): Cerata long, slender, flattened, smooth. Ceratal apex blunt. Digestive gland extending all along cerata through central duct; central duct bifurcating into perpendicular secondary branches terminating in bilateral and broad branches. Upon examination of translucent ceratal tissue, the _ uniformity and extent of the branching is visible. Buccal armature (Fig. 7B): Radular formula 26 X 0.1.0 (CASIZ 110358). Tooth wide and long, base extending anteriorly, covering posterior portion of front tooth. Cusp long and narrow in anterior portion of the tooth. Denticles visible on central part of each tooth, short and separate from each other, terminates in pointed tip; smooth masticatory border of jaw (Rudman, 1991). Reproductive system: Reproductive system similar to that described for the other species of Phyllodesmium, although, as in P. opalescens, the prostate gland is very large (Rudman, 1991). Phyllodesmium kabiranum Baba, 1991 (Figs. 1B, 8A—D) ?Eolida bella Ruppell & Leuckart, 1831: 35, Pl. 1. 10, fig. 4. ?Phyllodesmium bellum — O’Donoghue, 1929: 715. Phyllodesmium kabiranum Baba, 1991b: 113, figs. 4-5, Pie ties 2 Material examined: One specimen, CASIZ 89035, Seragaki Beach, 1.3 km ENE of Maeki-zaki, Okinawa, Page 14 The Veliger, Vol. 50, No. 1 Figure 7. Phyllodesmium colemani. A. View of digestive tract inside a ceras. B. Lateral view of radula. Abbreviations: dt, digestive tract. Scale bars: A = 5 mm; B = 0.02 mm. Ryukyu Islands, Japan, 1.5 m depth, 3 April 1993, R.F. Bolland. Two specimens, one dissected, CASIZ 103793, Cathedral Rock, Balayan Bay, Batangas Province, Luzon Island, Philippines, no depth, 25 February 1995, D.W. Behrens. Distribution: Known from its type locality Okinawa, Japan (Baba, 1991b) and the Philippines (present study). External morphology (Fig. 1B): Body 56 mm in length, large in size, narrowing uniformly from anterior to posterior end, wide foot, smooth and moderately long rhinophores, oral tentacles shorter in size than rhino- phores. Cerata flattened, smooth, with a straight apex, extending outwardly covering the whole dorsum. Ceratal arrangement of one precardiac arch and single vertical rows on each side of the body, lying on distinctive ridges, up to 7 to 8 cerata on each precardiac arch on each side of the body, 8 to 11 cerata on each of the 7 to 9 postcardiac single arches on each side dorsum. Reproductive opening found in right side of dorsum, right below and inside precardiac arch. Renal ope is in interhepatic space, right above the most basal po dge of the first postcardiac arch. Anal papilla on distinctive mound, between first and second postcardiac arches on right side of body. Cerata and digestive gland branching (Fig. 8C): Cerata flattened, smooth, terminating in a curled apex. Digestive gland branching into secondary and tertiary branches; numerous multiple branches that extend in “web-like’> manner, terminating in close proximity to the body wall. Ducts terminate in small chambers, capable of harboring zooxanthellae. Dark brownish- green color present likely due to presence of zooxan- thellae in the cerata (Baba, 1991b). Buccal armature (Figs. 8A—B): Radular formula 63 X 0.1.0 (CASIZ 89035). Base of tooth short and wide, long cusp. Denticles visible along center of tooth, separated, short, with blunt tip; masticatory border of the jaw plates has large denticles. Reproductive system (Fig. 8D): Preampulla duct nar- row, long; expanding into large, bulbous, folded ampulla. Ampulla divides into receptaculum seminis and penial complex by thin duct. Penial complex large, connecting to massive and folded albumen gland, opening to a folded, long prostate. Prostate connecting D. M. Ortiz & T. M. Gosliner, 2007 Page 15 Figure 8. Phyllodesmium kabiranum. A. Lateral view of radula. B. View of radular denticles. C. View of digestive tract inside a ceras. D. Reproductive system. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 0.12 mm; B = 0.05 mm; C = 12 mm; D = 5 mm. to massive, folded, rectangular shaped female gland mass. Female duct opening into a vagina at base of first cereal cluster on right side of dorsum. Remarks: Baba (1991) considered Eolida bella Ruppel & Leuckart, 1831 as a possible senior synonym of Phyllodesmium. Baba noted the similarity in color pattern between the two taxa. However, the remainder of the anatomy of E. bella remains unknown. Examination of material from the Red Sea is necessary to confirm the identity of E. bella, in order resolve this nomenclatural issue. Phyllodesmium macphersonae (Burn, 1962) (Figs. 1D, 9A—D) Cratena macphersonae Burn, 1962: 118-119, figs. 19— 20. Phyllodesmium macphersonae — Rudman, 1981b: 239— 242, 244-261, figs. 12A, B, 17, 21C, 27. Material examined: One specimen, dissected, CASIZ 115724, Horseshoe Cliffs, Okinawa, Ryukyu Islands, Japan, 3.0 m depth, 29 May 1998, R.F. Bolland. One specimen, CASIZ 104700, 14 km W of Ikei-shima, Tengan Pier, Okinawa, Ryukyu Islands, Japan, 2.1 m depth, 26 August 1994, R.F. Bolland. Distribution: Originally described from Victoria, Aus- tralia (Burn, 1962). Recorded from the Coral Sea (Lord Howe Island) and Tasmania, Australia (Rudman, 1991). Also collected from Japan (present study). External morphology (Fig. 1D): Body 23 mm in length, of moderate size, extending narrowly from the anterior to the posterior end. Anterior end of foot angular, The Veliger, Vol. 50, No. 1 Figure 9. Phyllodesmium macphersonae. A. Lateral view of radula. B. Close up view of radular denticles. C. View of digestive tract inside a ceras. D. Reproductive system. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 0.04 mm; B = 0.02 mm; C = 10 mm; D = 5 mm. narrow, elongate, tapering from the anterior to posterior end. Rhinophores moderately long, smooth, close in size to oral tentacles. Ceratal arrangement of one precardiac arch, up to 6 to 8 single vertical postcardiac rows, up to 6 to 8 cerata on each postcardiac rows on each side of body. Reproductive opening on right side, below anterior basal edge of precardiac arch. Renal opening immediately above first postcardiac row, in interhepatic space. Anal papilla between first and second postcardiac row, immediately below uppermost postcardiac row. Cerata and digestive gland (Fig. 9C): Cerata cylindri- cal, smooth, terminating in curled apex. Digestive D. M. Ortiz & T. M. Gosliner, 2007 Page 17 Figure 10. Phyllodesmium briareum. A. View of digestive tract inside a ceras. B. Lateral view of radula. C. Reproductive system. Abbreviations: a, albumen gland; am, ampulla; rs, receptaculum seminis; pc, penial complex; fmg, female mass gland; dt, digestive tract. Scale bars: A = 10 mm; B = 0.01 mm; C = 5 mm. gland composed of central duct running along entire ceras. Photographs of the cerata do not obviously show the detailed structure. However, under close examina- tion through the dissecting microscope, the arrange- ment of the secondary and tertiary branches was similar to that of P. magnum. Buccal armature (Fig. 9A—B): Radular formula 28 X 0.1.0 (CASIZ 65346). Tooth long and narrow base, long cusp. Denticles visible, all along central region of radular teeth, separated, short, terminating in a blunt tip. Masticatory border of jaw smooth (Baba, 1991). Reproductive system (Fig. 9D): Preampulla narrow, reduced, duct expanding into long and thin ampulla by ‘irregular narrow duct; duct bifurcating into receptacu- lum seminis and penial complex. Penial complex extending into small and bulbous prostate, which connects to a long, folding and thin female gland mass; female duct opened to a vagina at the base of first ceratal cluster. Phyllodesmium briareum (Bergh, 1896) (Figs. 1G, 10A—C) Ennoia briareus Bergh, 1896: 392-4, Pl. 16, figs. 14, 15. Phyllodesmium briareus — Gosliner et al., 1996: 177, fig. 627. Phyllodesmium briareum — Rudman, 1991: 181-187, figs. 13, 14A,B, 15, 16, 17, 27-29. Material examined: Twenty specimens, CASIZ 065346, Barracuda Point, W side “Pig Island,’ near Madang, north coast, Papua New Guinea, 15.2 m depth, 13 January 1988, T.M. Gosliner. One specimen, one dissected, CASIZ 065299, Madang (near lighthouse), north coast, Papua New Guinea, 7.6m depth, 22 January 1988, T.M. Gosliner. Four specimens, one dissected, CASIZ 83678, Devil’s Point (SW side of Maricaban Island), Maricaban Island, Batangas Prov- ince, Luzon, Philippines, 19 February 1992, T.M. Gosliner. Page 18 Distribution: Found in the Philippines, Malaysia and Papua New Guinea (Rudman, 1991). Recent studies found P. briareum in Japan and Indonesia (present study). External morphology (Fig. 1G): Body 16 mm in length, moderate size, narrow, reduced (in some cases more elongate). Foot narrow, angular anterior end of corners. Oral tentacles slightly longer than greatly extended rhinophores. Smooth, cylindrical cerata terminating in blunt tip. Ceratal arrangement of one single-rowed precardiac arch, consisting 6 to 8 ceras on each side of body. Reproductive system, renal opening and anal papilla located on right side of body, underneath the precardiac row. Renal opening in the interhepatic space. Anal papilla below first post-cardiac row.. Postcardiac cerata in single rows across dorsum on each side of body, each side with 6 to 8 postcardiac clusters with up to 7 ceras on each of them. Cerata and digestive gland (Fig. 10A): Cerata long, slender, smooth, cylindrical, with branched digestive tract. Digestive tract branching into simple branches, in turn extending into secondary and tertiary branches, making for a dense layer and expansion of the digestive tract all across cerata. Buccal armature (Fig. JOB): Radular formula 34 xX 0.1.0 (CASIZ 83678). Base of tooth wide, long, with long cusp. Denticles visible, along central edge of each tooth, short, well-separated, terminating in pointed radular tip. Masticatory border of jaw smooth (Rud- man, 1991). Reproductive system (Fig. JOC): Preampullary duct short, narrow, expanding into bulbous, long and folded ampulla. Ampulla connecting through broad duct to receptaculum seminis and penial complex. Penial complex opening to wide and short prostate through broad duct. Duct connecting to folded albumen gland, located on massive membrane gland. Prostate connect- ing into folded and long female gland mass. Phyllodesmium longicirrum (Bergh, 1905) Myrrhine longicirra Bergh, 1905: 227-9, P1. 20, figs. 20-29. Phyllodesmium longicirra — Rudman, 1981b: 242-261, figs. 16, 18-20, 24-27. Phyllodesmium longicirrum — Rudman, 1991: 195, figs. 26, 28, 29. Distribution: Described originally from Indonesia (Bergh, 1905), also recorded from the Great Barrier Reef (Rudman, 1991). Discussion: The anatomy of this species has been lescribed by Rudman (1981b; 1991). The Veliger, Vol. 50, No. 1 Phyllodesmium pecten Rudman, 1981 Phyllodesmium pecten Rudman, 1981b: 237-239. figs. 4A IS) 16@) Ds Es 27- Distribution: Known only from its type locality Dar es Salaam, Tanzania (Rudman, 1981b). Discussion: Anatomy of species described by Rudman, 1981b. Phyllodesmium opalescens Rudman, 1991 Phyllodesmium opalescens Rudman, 1991: 177-181, figs. 9, 10, 11, 12, 28, 29. Distribution: Known only from its type locality Hong Kong (Rudman, 1991). Discussion: Anatomy of species described by Rudman, 1991. Phyllodesmium horridum (Macnae, 1954) Favorinus horridus Macnae, 1954: 19-21, figs. 11-13, Pl. 1, fig. 4. Phyllodesmiopsis horridus — Risso-Dominguez, 1964: 222-238. Phyllodesmium horridus — Rudman, 1981b: 224. Phyllodesmium horridum — Gosliner, 1987: 124, fig. DIPIS) Distribution: Known and described originally from South Africa (Macnae, 1954). Discussion: Anatomy of species described by Rudman (1981b; 1991). Phyllodesmium iriomotense Baba, 1991 Phyllodesmium iriomotense Baba, 1991b: 115, figs. 6-7, Bilis hee 3 Distribution: Known only from its type locality Okinawa, Japan (Baba, 1991b). Discussion: Anatomy of species described by Baba, 199 1b. Phyllodesmium guamensis Avila et al., 1998 Phyllodesmium guamensis Avila et al., 1998: 148, figs. 1-10. Distribution: Only found in its type locality Guam (Micronesia) (Avila et al., 1998). Discussion: Anatomy of species described by Avila et al., 1998 D. M. Ortiz & T. M. Gosliner, 2007 Phyllodesmium jakobsenae Burghardt & Wagele, 2004 Phyllodesmium jakobsenae Burghardt & Wagele, 2004: 1, figs. 1—S. Distribution: Known only from Indonesia (Burghardt & Wagele, 2004). Discussion: The anatomy of this species was completely described by Burghardt & Wagele (2004). Phyllodesmium rudmani Burghardt & Gosliner, 2006 Phyllodesmium jakobsenae Burghardt & Gosliner, 2006: 31, figs. 1-5. Distribution: Known only from Indonesia (Burghardt & Gosliner, 2006). Discussion: The anatomy of this species was completely described by Burghardt & Gosliner (2006). Godiva quadricolor (Barnard, 1927) Hervia quadricolor Barnard, 1927: 203, Pl. 20, figs. 9, 10. Godiva quadricolor — Macnae, 1954: 23-25, text figs. 14-16. Distribution: This species has been collected from South Africa (Barnard, 1927) and Western Australia (Willan, 1987). Discussion: Anatomy of species described by Willan (1987). Godiva Macnae has been placed on the Official List of Generic Names in Zoology with the name number 1717 (I.C.Z.N, 1966, Opinion 778). The name quadricolor Barnard, as published in the binomen Hervia quadricolor (type species of Godiva Macnae), has been placed in the Official List of Specific Names in Zoology with name number 2148 (1.C.Z.N., 1966, Opinion 778). Favorinus japonicus Baba, 1949 Favorinus japonicus Baba, 1949: 177, Pl. 43, fig. 151, figs. 135-136. Distribution: Have been found in Dar es Salaam, Tanzania (Rudman, 1980) and Japan (Baba, 1949) and throughout the Indo-Pacific, extending from the Western Indian Ocean to the Hawaiian Islands (Gosliner, 1980). Discussion: Anatomy of species described by Baba (1949) and Rudman (1980). Page 19 RESULTS We performed several analyses of the data matrix (Table 3). After several iterations we re-examined the characters. Three characters (14, 21 and 30) were then deleted (using PAUP 4.0 and manually) from the analysis because they are continuous and it 1s difficult to assess discrete character states. This analysis yielded two trees of 64 steps and consistency and retention indices of 0.469 and 0.730 respectively (Fig. 11). From this analysis, the monophyly of Phyllodesmium 1s supported. Our strict consensus tree shows Phyllodes- mium as a monophyletic group, supported by a Bremer value of 2 and defined by four synapomorphies: character 12, angular foot corners; character 15, rhinophores smooth or slightly nodular; character 16, masticatory border with a single row of denticles; character 27, cnidosac absent (Fig. 12). Most nodes have a Bremer support value of 1. The third node above the basal node has a value of 3. Additionally, the clade that contains P. poindimiei, P. briareum, P. macphersonae and P. colemani, the clade that contains P. magnum, P. longicirrum, P. guamensis, P. paran- gatum, P. jakobsenae and P. rudmani and the clade containing P. hyalinum and P. crypticum each have a value of 2. Our phylogenetic analysis shows that species pos- sessing an unbranched digestive tract (P. horridum, and P. opalescens) are most basal, while others that have a branched digestive tract are more derived. Phyllodes- mium species having a branched digestive tract are included in one clade, indicating that elaboration of the digestive gland duct represents a single evolutionary event. Members of this clade share other common traits relating to jaw morphology, ceratal structure, and their ability to store zooxanthellae. Within this group, there is one well-supported clade that divides into two sister clades. The first one 1s supported by one synapomor- phy (narrow foot), while the second is supported by two synapomorphies (ceratal apex curled and tightly congested radular denticles). Based on the present phylogenetic analysis, some characters used in this study exhibit at least one instance of reversal (indicated by an underline of the character number in Figure 12). Even though the instances of homoplasy are moderate, the phylogenetic reconstruction of the ancestral state is unambiguous. Figure 12 shows some of the characters displaying homoplasy, such as the presence of character 2 in several taxa within different lineages. Several derived characters support distinct phylogenetic relationships within various subclades of Phyllodesmium. These derived characters include the presence of zooxanthel- lae, ceratal morphology (apex, surface, shape) and ceratal arrangement, digestive tract branching, and masticatory border of jaw. Tracing of character trace Page 20 Favorinus japonicus Godiva quadricolor P. serratum P. poindimiei P. briareum P. macphersonae P. colemani P. iriomotense P. horridum Figure 11. evolution was made on the not fully resolved consensus tree rather than on one of the two fully resolved trees produced by the phylogenetic analysis. This approach emphasizes the distribution of characters that are consistent rather than those that vary. DISCUSSION The phylogenetic analysis carried out in this paper supports the monophyly of the exclusively tropical Indo-Pacific species that have been included in Phyllodesmium. In general, the non-parsimony based scenario of evolution suggested by Rudman (1991) is generally upheld by the present phylogenetic analysis. More specifically, our parsimony-based phylogeny Rudman’s view that more basal members have relatively simple digestive gland branching and more derived taxa have more complex branching of upports P. magnum The Veliger, Vol. 50, No. 1 ~ = y Pa SS es S =e x 2 8 oes = = s D BSS SS So Sale o SN) a 3 > S SES aH = ~ S S D Sos OS v SO S S 9 Ww — SS a — s 3s S 3s 8 © Sawa = © 2 S 8 ff SS AL A AL RR CR RE RS ee Strict consensus phylogeny of Phyllodesmium, (characters 14, 21, 30 excluded). digestive gland ducts to provide greater surface area for photosynthesis in species that harbor zooxanthellae. However, the arrangement of taxa within Rudman’s branching diagram differs in some respects from our phylogeny. For example, Rudman suggested that P. serratum was most basal within Phyllodesmium, while our analysis suggests that P. opalescens is more basal. Additionally, Rudman suggested a continuum of evolution from P. crypticum to P. hyalinum to P. pecten, whereas our analysis suggests that P. crypticum and P. hyalinum are sister species, while P. pecten is a member of another subclade. Rudman treated evolution within Phyllodesmium based on the comparative examination of the digestive system and cerata. However, it is hard to assess how related these assumptions are due to insufficient data regarding the specificity of the feeding specialization across all Phyllodesmium taxa. It is evident, though, D. M. Ortiz & T. M. Gosliner, 2007 Bo oeoge LL N A NY) 3 a) S nw as Ss fos oS 2 £ cs SS SS eSo Sa aes = eS USS SS 8 5 Ss SSS Os wees 8 5 = 8 & R=) A) ek aiee Sal Sone Qn 2 am A OD D2, iQ 24 aS) 1 5 2Y 3 12 8,16,25+ 2 18 D 23 1 JUL BS “i 20 24 1 16 1 28 29 Figure 12. Page 21 Sue S S = en eS & & 3 & & Sew Sas S & SS SS © 8 Ss 6-8 S 8 of BO OSES See ese aN SS 6 & S&S § a & SS AR CINE Rh | ay rei Westen Guerin (a Ta 2 zo) 13 Ria ce wo 24 Iti |2 oe 4,23 17/6 1 DO ars 22 94178 ve 1 +.6.20 2 1+29./0 1+ 25 D Bele 57326 18,22,24 9) = SID IS NO 2H. Character and Bremer support analysis of strict consensus tree. Italicized numbers refer to characters contained in text. Underlined numbers indicate instances of reversals. Bold numbers are Bremer support values. that the morphological changes occurring within Phyllodesmium.are correlated with their feeding behav- ior. The increase in size, and the flattening and branching of the digestive tract are all characteristics of the cerata that have developed across taxa of Phyllodesmium. These characteristics allow a more efficient means of transporting and accommodating the zooxanthellae, and displaying mimicry of the octocoral coelenterates they prey upon (Rudman, 1981b, 1991). Histological and digestive dissections on specimens living or freshly preserved material will be needed to assess the ecological interaction occurring across the Phyliodesmium taxa. Specimens, used in this study, were fixed in formalin or Bouin’s solution making any presence of calcium carbonate octocoral sclerites undetectable and unidentifiable by the observer. Hence, fresh material is needed in order to assess the feeding sources and ecological interactions occurring between the nudibranch species and its octocoral prey. Further descriptions of Phyllodesmium species will help to resolve the phylogenetic relationships and symbiotic relationships within Phyllodesmium. In addition, mo- lecular data would aid in providing more information on the evolutionary trend of this genus. Combining these methods would provide a clearer view on the evolution of Phyllodesmium and its relationship with its symbionts. Acknowledgments. This paper was supported by San Francisco State University, through its NIH predoctoral fellowship program. This work was also supported by a National Science Foundation PEET grant DEB 0329054, Phylogeny of the Nudibranchia, awarded to the second author and Angel Valdés. This paper is a manuscript of the senior author’s Masters thesis. The authors thank Robert Bolland and Dr. William Rudman for their contributions of specimens and photographs used in this study. Also thanks to Dr. Gary Williams, Dr. Angel Valdés, Erin Rempala, Elizabeth Ruck and Yvonne Valles in the Invertebrate Zoology and Geology Department of the California Academy of Sciences, and Dr. Thomas Niesen at San Francisco State University for their guidance, support and comments through the preparation of this manuscript. Above all, the senior author would like to thank her family and friends for their patience, love and encouragement. LITERATURE CITED AVILA, C., M. BALLESTEROS, M. SLATTERY & V. J. PAUL. 1998. 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Sinauer Associates: Sunderland, Massachusetts. WILLAN, R. C. & N. COLEMAN. 1984. Nudibranchs of Australasia, 56 pp, Australasian Marine Photographic Index: Sydney. WILLAN, R. C. 1987. Phylogenetic systematics and zoogeog- raphy of Australian nudibranchs. 1. Presence of the aeolid Godiva quadricolor (Barnard) in Western Australia. Journal of the Malacological Society of Australia 8:71—85. The Veliger 50(1):24-26 (March 11, 2008) THE VELIGER © CMS, Inc., 2007 Earliest Record of the Genus Haliotis (Mollusca: Gastropoda) from the Late Cretaceous (Campanian) of Los Angeles County, California LINDSEY T. GROVES Natural History Museum of Los Angeles County, Malacology Section, 900 Exposition Boulevard, Los Angeles, CA 90007 (e-mail: lgroves@nhm.org) JOHN M. ALDERSON Natural History Museum of Los Angeles County, Invertebrate Paleontology Section, Research Associate, 900 Exposition Boulevard, Los Angeles, CA 90007 (e-mail: jjalderson@earthlink.net) Abstract. Cretaceous abalone are extremely rare and are known from only two valid species: Haliotis lomaensis Anderson, 1902, from the Late Cretaceous (latest Campanian/earliest Maastrichtian) of San Diego County, California and H. antillesensis Sohl, 1992, from the Late Cretaceous (late Maastrichtian) of southwestern Puerto Rico. The earliest record of the genus Haliotis is here documented from Late Cretaceous (middle middle to late middle Campanian) strata of the Tuna Canyon Formation, Garapito Creek area of Topanga Canyon, Santa Monica Mountains, Los Angeles County, California. This additional Cretaceous record for Haliotis could possibly indicate a North American origin for the family Haliotidae. INTRODUCTION Cretaceous abalone species are extremely rare, current- ly comprising only two valid species (Sohl, 1992; Geiger & Groves, 1999; Geiger, 2000). These species include: Haliotis Anderson, 1902 from the Late Cretaceous (latest Campanian/earliest Maastrichtian) Point Loma Formation, San Diego County, California and H. antillesensis Sohl, 1992 from the Late Creta- ceous (late Maastrichtian) El Rayo Formation, Sabana Grande quadrangle, southwest Puerto Rico. Suspect records of Cretaceous abalone that have been relegated to pleurotomarioidean genera include Haleotis? [sic] antiqua Binkhorst, 1861 [ Trochus limburgensis Kaunhowen, 1897] from the Maastrichtian near Maastricht, Limburg Province, the Netherlands, Pleur- otomaria antiqua (Binkhorst, 1861) of Weinzettl (1910) from the Late Cretaceous (Cenomanian) near Kor- yeany, Czech Republic, and Haliotis cretacea Lundg- ren, 1894 [= Pleurotomaria sp.?] from the Late Cretaceous (Campanian) near Barnakallegrottan, southeastern Sweden [see Sohl (1992) for additional details]. A poorly preserved specimen identified as Haliotis sp. by Dawson (1978) and reported by Sundberg (1979, 1984) from the Late Cretaceous (Maastrichtian) Cabrillo Formation of San Diego County, California is actually a specimen of the calyptraeid genus Lysis (L.R. Saul, personal commu- nical lomaensis i0n). MATERIAL A single poorly preserved internal mold with little remaining original or recrystallized shell material, Natural History Museum of Los Angeles County, Invertebrate Paleontology (LACMIP) hypotype 13237 (Figs. 1-4) from LACMIP loc. 27110 (ex University of California, Los Angeles loc. 7110) that measures 5.9mm in overall length retains several diagnostic features that validate it as a haliotid. Observed features include a flattened “shell” with low spire, wide columella, and a row of six tremata toward the left periphery. Because of such poor preservation we hesitate to describe a new species based on this sole example. Nevertheless, due to the uniqueness of this specimen from Late Cretaceous (middle middle to late middle Campanian) strata, it is noteworthy enough to mention as the earliest known worldwide representative of the genus Haliotis. LOCALITY LACMIP loc. 27110 is on the north side of Garapito Creek just above the 1300 ft. contour line, 900 ft. north, 735 ft. east of SW corner of section 33 (projected), TIN, RI6W San Vicente y Santa Monica Land Grant, northeast of Sylvia Park, United States Geological Survey (USGS) Topanga quadrangle (1976 ed.), Santa Monica Mountains, Los Angeles County, California. L. T. Groves & J. M. Alderson, 2007 Figures 1-4. AHaliotis sp., hypotype LACMIP 13237, from LACMIP loc. 27110, 1 = dorsal view (7.6), 2 = oblique view (X10), 3 = lateral view (<9.3), 4 = columellar view (10.3). STRATIGRAPHY & AGE The specimen was collected by the junior author on 21 December, 1983 from the base of informal ““member D” of Wilson (1941) [= map unit Ktd of Yerkes et al. (1994)] within the Late Cretaceous (late middle to early late Campanian), Metaplacenticeras pacificum ammon- ite zone [33N chron] (Elder & Saul, 1996) part of the Tuna Canyon Formation of Yerkes & Campbell (1979) [= Unnamed strata of Dibblee, 1992]. This ammonite zone was cited as Late Cretaceous (middle middle to late middle Campanian) [C33 chron] by Squires & Saul (2003) and we follow this usage. ““Member D” is a fossiliferous fine-grained sandstone that occurs imme- diately above a thick, cobble conglomerate informally designated as “member C” by Wilson (1941) [= map unit Ktc of Yerkes et al. (1994)] and is equivalent to the lowermost part of a fine-grained sandstone reported by Popenoe (1954). LACMIP loc. 27110 is within an unusual small lens of “member D” that disappears along strike within 100 ft. (33 m). Unfortunately, Dibblee (1992) incorrectly mapped this lens of ““member D” as Paleocene Santa Susana Formation as did Yerkes et al. (1994). However, recent field work by the junior author combined with the fauna listed below, correctly place the locality within the Tuna Canyon Formation. In addition to the ammonite Metaplacenticeras pacificum (Smith, 1900), LACMIP 27110 also yielded the ammonite Baculites cf. B. inornatus Meek, 1862, the gastropods Aftira sp., Turritella chicoensis pescaderoen- sis Arnold, 1908, Gyrodes pacificum Popenoe & others, 1987, Volutoderma n. sp., and Biplica obliqua (Gabb, 1864), and the bivalves Prerotrigonia evansana (Meek, 1858), Glycymeris veatchii (Gabb, 1864), mytilid sp., Ostrea sp., and Calva sp. PALEOBIOGEOGRAPHY Geiger & Groves (1999), Geiger (2000), and Geiger & Poppe (2000) discussed three possible haliotid radiation Page 25 models as follows: 1) An “Indo-Pacific” model, also discussed by Lindberg (1992), indicates that living abalone are most diverse in the central Indo-Pacific, which implied that this was their center of radiation; 2) A “Pacific Rim” model proposed by Talmadge (1963), where abalones originated on an island arc from Japan to northern Australia and radiated to California, southern Australia, and the Indo-Pacific; and 3) A “chromosomal” model where species with a low diploid number (28) live in the eastern Mediterranean Sea and species with higher diploid numbers (32) in the Indo-Pacific and (36) in the North Pacific, abalones dispersed eastward from the Mediterranean. However, because these models do not consider the fossil record they could be rejected. Moreover, this confirmation of the earliest known haliotid from Late Cretaceous (middle middle to late middle Campanian) strata of southern California combined with the fact that Cretaceous abalones are known exclusively from North America further strengthens the possibility of a North American origin for the family Haliotidae. Kiel & Bandel (2000) described Temnotropis frydai (Family Temnotropidae) from the Late Creatceous (late Cam- panian) Valcarga Formation near Torallola, Lérida Province, Catalona Region, northeastern Spain and speculated that Temmnotropis is a likely ancestor of Haliotis. Temnotropis has a Haliotis-like shell with a slit rather than a row of tremata. With a possible ancestor in Spain, the haliotids may have originated in the eastern Atlantic (S. Kiel, personal communication, 2007). However, should a haliotid be found in strata older than middle middle to late middle Campanian a reevaluation of their origin will be necessary. Acknowledgments. We express our thanks to our colleagues Richard L. Squires (California State University, Northridge, Geological Sciences), LouElla R. Saul (LACMIP), and Angel Valdes (LACM Malacology) for reviewing the manuscript and adding valuable suggestions. Daniel L. Geiger (Santa Barbara Museum of Natural History) is acknowledged for examining the specimen, confirming the identification, and reviewing the manuscript. His unsurpassed knowledge of abalone morphol- ogy and phylogenetics is greatly appreciated. Many thanks to Steffen Kiel (University of Leeds, Leeds, England, UK) for his thoughtful review of the manuscript and valuable insights. LouElla R. Saul identified additional mollusks from LACMIP loc. 27110. Special thanks to N. Scott Rugh (San Diego Natural History Museum) for the loan of Dawson’s (1978) Haliotis sp. Angel Valdés is also thanked for assisting with digital photography. Many thanks to Cathy L. Groves (LACM Echinoderms Section) for assistance with digital image manipulations. PERE RATUINE, Cie D ANDERSON, F. M. 1902. Cretaceous deposits of the Pacific Coast. Proceedings of the California Academy of Sciences, 3rd ser., Geology 2(1):1—154, pls. 1-12. ARNOLD, R. 1908. Descriptions of new Cretaceous and Tertiary fossils from the Santa Cruz Mountains, Califor- nia. Proceedings of the United States National Museum 34(1617):345-390, pls. 31-37. BINKHORST, J.-T. 1861. Monographie des Gastéropodes et des Céphalopodes de la Craie Supérieure du Limbourg. C. Muquardt: Bruxelles, Belgique. 83 pp, [gastropods] + 44 p. [cephalopods]. Dawson, M. K. 1978. The paleontology of the Cabrillo Formation [M.S. thesis]. San Diego State University: San Diego, California. 90 pp, 7 figs, 2 pls. DIBBLEE, T. W., JR. 1992. Geologic map of the Topanga and Canoga Park (south '%) quadrangles, Los Angeles County, California. Dibblee Geological Foundation Map DF-35, | sheet, 1:24,000. ELDER, W. P. & L. R. SAUL. 1996. Taxonomy and biostratigraphy of Coniacian through Maastrichtian Anchura (Gastropoda: aporrhatdae) of the North Amer- ican Pacific Slope. Journal of Paleontology 70(3):381—399, figs. 1-6. GABB, W. M. 1864. Description of the Cretaceous fossils. Geological Survey of California, Palaeontology 1(4):57— 217, pls. 9-32. : GEIGER, D. L. 2000. Distribution and biogeography of the Recent Haliotidae (Gastropoda: vetigastropoda) world- wide. Bollettino Malacologico 35(5—12):57—120, figs. 1— 192. GEIGER, D. L. & L. T. GRoves. 1999. Review of fossil abalone (Gastropoda: vetigastropoda: haliotidae) with comparison to Recent species. Journal of Paleontology 73(5):872-885, figs. 1-3. GEIGER, D. L. & G. T. PopPE. 2000. The family Haliotidae. A Conchological Iconography. ConchBooks: Hackenheim, Germany. 135 pp, 80 figs. + numerous unnumbered figs., 83 pls. KAUNHOWEN, F. 1897. Die Gastropoden der Maestrichter Kreide. Palaeontologische Abhandlungen herausgegben von W. Dames und E. Koken, neue folge 4(1):3—126, figs. 1-34, pls. 1-54. KIEL, S. & K. BANDEL. 2000. New slit-bearing Archaeogas- tropoda from the Late Cretaceous of Spain. Berliner geowissenschaftliche Abhandlungen, Reihe E 34:269-277, pl. 1. LINDBERG, D. R. 1992. Evolution, distribution, and system- atics of Haliotidae. In: S. A. Shepard, M. J. Tegner & S. A. Guzman del Proo (eds.), Abalone of the world: Biology, fisheries and culture. Fishing News Books: Oxford, England, Chapter 1, p. 3-18, figs. 1.1—1.5. LUNDGREN, B. 1894. Jamf6relse mellan Molluskfaunan I Mammillatus och Mucronata Zonera I Nordéstra Skane (Kruistianstadsomradet). Svenska Vetenskaps-akade- miens Handlingar ser. 4 26:3—58, pls. 1-2. MEEK, F. B. 1858. Descriptions of new organic remains from The Veliger, Vol. 50, No. 1 the Cretaceous rocks of Vancouver Island. Transactions of the Albany Institute 4:37-49. MEEK, F. B. 1862. Descriptions of new Cretaceous fossils collected by the North-Western Boundary Commission, on Vancouver and Sucia islands. Proceedings of the Academy of Natural Sciences of Philadelphia 13:314-318. POPENOE, W. P. 1954. Mesozoic formations and faunas, southern California and northern Baja California. In: R. H. Jahns (ed.), Geology of southern California. California Division of Mines Bulletin 170:Pp. 15-21 figs. 1-4. Chapter 3, Historical Geology. POPENOE, W. P., L. R. SAUL & T. SUSUKI. 1987. Gyrodiform gastropods from the Pacific coast Cretaceous and Paleocene. Journal of Paleontology 61(1):70—100, figs. 1-7. SMITH, J. P. 1900. The development and phylogeny of Placenticeras. Proceedings of the California Academy of Sciences, 3rd ser., Geology 1(7):180—240, pls. 24-28. SOHL, N. F. 1992. Upper Cretaceous gastropods (Fissurelli- dae, Haliotidae, Scissurellidae) from Puerto Rico and Jamaica. Journal of Paleontology 66(3):414-435, figs. 1— 10. SQUIRES, R. L. & L. R. SAUL. 2003. New Late Cretaceous epitoniid and zygopleurid gastropods from the Pacific slope of North America. The Veliger 46(1):20—49, figs. 1— 71. SUNDBERG, F. A. 1979. Upper Cretaceous macro-fossils of San Diego. In: P. L. Abbott (ed.), Geological excursions in the southern California area. Department of Geological Sciences Publication, San Diego State University, Pp. 173— 176, figs. 1-2. pl. 1 (also published in 1984 with identical title In: Abbott, P.L. (ed.), Upper Cretaceous depositional systems, southern California — Northern Baja California. Pacific Section, Society of Economic Paleontologists and Mineralogists Guidebook 36:37—-40, figs. 1-2, pl. 1.) TALMADGE, R. R. 1963. Insular haliotids in the western Pacific. The Veliger 5(3):129-139, pl. 14. WEINZETTL, V. 1910. Gastropoda Ceského Kridového Utvaru. Palaeontographica Bohemiae 8:1—56, pls. 1—7. WILSON, H. D. B. 1941. Stratigraphy of the Cretaceous and Eocene rocks of the Santa Monica Mountains [Ph.D. minor dissertation]. California Institute of Technology: Pasadena, California. II + 31 pp, 4 figs., 3 pls. YERKES, R. F. & R. H. CAMPBELL. 1979. Stratigraphic nomenclature of the central Santa Monica mountains, Los Angeles County, California. United States Geological Survey Bulletin 1457-E:iv + El E31, figs. 1-5, pls. 1-3. YERKES, R. F., R. H. CAMPBELL & J. M. ALDERSON. 1994. Preliminary geologic map of the Topanga quadrangle, southern California. United States Geological Survey Open File Report 94-266:2—14, pls. 1-2. The Veliger 50(1):27—30 (March 11, 2008) THE VELIGER © CMS, Inc., 2007 Predatory Behavior and Diet of Eupleura sulcidentata Dall, 1890 (Gastropoda: Muricidae) from West Florida GREGORY S. HERBERT AND SHUBHABRATA PAUL Department of Geology, University of South Florida, Tampa, FL 33620, USA (e-mail: gherbert@cas.usf.edu) Abstract. The diet and feeding behavior of the muricid gastropod Eupleura sulcidentata are documented for the first time from laboratory aquarium experiments. Eupleura sulcidentata feeds readily on a broad range of shelled invertebrate prey, including barnacles, bivalves and calyptraeid gastropods, by drilling. Drillholes are small (<1 mm) but tend to have beveled sides, a morphology generally regarded as diagnostic of naticid drillholes. Key Words: Gastropoda, Muricidae, Eup/eura, predation, diet. The present study contributes new observations on the diet and feeding behaviors of Eupleura sulcidentata Dall, 1890, a diminutive species of ocenebrine muricid with a range that extends from Florida to the Bahamas and northern Cuba. Previously, the feeding biology of this species was studied by Radwin and Wells (1968), who found £. sulcidentata difficult to maintain in captivity. In their experiments, E. sulcidentata preda- tors were offered several types of invertebrate prey, including the bivalves Crassostrea virginica, Ostrea equestris, and Brachidontes exustus, and two barnacles, but refused all five and eventually died without feeding. Predatory gastropods of the family Muricidae are well known for their capacity to drill holes in shelled invertebrate prey (Carriker, 1961, 1981; Carriker and Gruber, 1999; Carriker et al., 1974), but many muricids employ additional (or alternative) modes of food aquisition, including use of anesthetizing toxins (West et al., 1994; Roller et al., 1995), mechanical shell breaking and wedging (Wells, 1958; Dunkin and Hughes, 1984; Perry, 1985), ovophagy (Philipps, 1969; Taylor, 1976; Abe, 1983), kleptoparasitism (Ishida, 2001, 2004), carrion feeding (Wu, 1965; Morton, 1994), and true parasitism (Ward, 1965; Robertson, 1970; Matsukuma, 1977). Given this diversity, it is possible that E. sulcidentata is an obligate non-driller specializ- ing on prey other than those provided by Radwin and Wells (1968). As an initial test of this hypothesis, we repeated the Radwin and Wells experiment using prey known to oceur in the same microhabitat as E. sulcidentata. At least one of the prey species offered in the Radwin and Wells study, the oyster C. virginica, lives in muddier, lower salinity waters of the upper estuary, whereas E. sulcidentata is found only in sandier, normal marine conditions of the lower estuary and shallow coast. Thus, in at least one case, E. sulcidentata may have simply been refusing an unfamiliar prey. Although E. sulcidentata is not uncommon, its cryptic microhabitat makes it difficult to observe its feeding preferences and behaviors in the field. In Tampa Bay, Florida, E. sulcidentata is most common in tidal channels between mangrove islands, where the channels are paved with the disarticulated valves of the venerid clam Mercenaria campechiensis (Gmelin, 1791). When oriented in a hydrodynamically-stable, concave- down position, large Mercenaria valves create a cave- like domicile for E. sulcidentata and numerous other small invertebrates. Because of their local abundance and close proximity to E. sulcidentata, these other invertebrates are the most likely components of the diet of E. sulcidentata. Predators and prey in this experiment were collected between December 2005 and February 2006 in 1—2 m of water from Miguel Bay, in the southernmost region of Tampa Bay, Florida and transferred to a laboratory at the University of South Florida in Tampa. Six E. sulcidentata, all roughly 20 mm in shell length (maxi- mum shell length of this species) and presumably mature, were collected from the field site, all of them underneath overturned Mercenaria valves. Nine shelled invertebrates encountered with E. sulcidentata preda- tors in this microhabitat were collected and offered as food in this experiment, including the barnacle Balanus eburneus Gould, 1841; five species of bivalve: Ostrea equestris Say, Brachidontes exustus (Linneaus, 1758), Timoclea grus (Holmes, 1858), Lyonsia floridana Conrad, 1849, and Anomalocardia auberiana (dOr- bigny, 1842); two species of slipper limpet: Bostryca- pulus aculeatus (Gmelin, 1791) and a member of the Page 28 ihe Veligers Vole s0 Nom Figure 1. Examples of predatory drillholes produced by the muricid gastropod Eupleura sulcidentata in barnacle, bivalve, and gastropod prey. Length measurements are for the anterior-posterior shell axis unless otherwise stated. A. Balanus eburneus (height 4.3 mm). B. Balanus eburneus (height 5.1 mm). C. Ostrea equestris (15.2 mm). D. Brachidontes exustis (5.9 mm). E. Bostrycapulus aculeatus (14.4 mm). F. Crepidula depressa (15.5 mm). G. Anomalocardia auberiana (13.2 mm). H. Lyonsia floridana (6.9 mm). I. Timoclea grus (7.3 mm). G. S. Herbert & S. Paul, 2007 Crepidula plana species complex, probably Crepidula depressa Say, 1822 (see Collin 2001), and the chiton Isnochiton papillosus (C. B. Adams, 1845). The six predators were housed in a single 10-gallon laboratory aquarium with recirculating seawater (changed weekly) from the bay. Seawater was main- tained at a constant salinity of 35 ppt and a temperature of 15-18°C to mimic conditions at the field site during the time of collection. Within the aquarium, predators were isolated from one another by placing each in its own 10 X 7 X 5 cm clear plastic box into which ten 0.5 cm diameter holes had been drilled. The holes provided ample water circulation, and the box is approximately the same volume as the cryptic microenvironment beneath disarticulated Mercenaria valves. The boxes also allowed us to observe the activities of individual predators and monitor the feeding experiments contin- uously. Prey were offered one species at a time to predator boxes, except for Crepidula and Bostrycapulus, which were collected on the same Mercenaria valve and offered simultaneously to a single predator. Cemented prey, such as oysters and barnacles, and sessile, loosely attached prey, such as Brachidontes and Crepidula, which were found attached to the interiors of Mercenaria shells, were introduced to the boxes on the original Mercenaria shell cut down to 5 cm? pieces. All dead prey shells were removed prior to intro- ducing the Mercenaria piece to the predator. Anom- alocardia, which is free-living and shallowly infaunal, was added to a box with sand 1 cm deep to allow natural burial and to determine whether the predator could excavate buried prey. Feeding experiments were monitored over a 2-month period, and predated shells were removed daily. Eupleura sulcidentata fed readily upon eight of the nine prey species offered, the exception being the highly mobile [snochiton papillosus, which was never attacked and experienced no mortality during the experiment. All other prey species were successfully drilled and eaten, in contrast to the results of the Radwin and Wells (1968) experiment. Eupleura sulcidentata preda- tors produced drillholes with a mean outer borehole diameter of 0.57 mm + 0.085 (n = 40) and, in most cases, beveled sides (i.e., a naticid-like morphology). Drillholes in Anomalocardia were more often straight- sided (i.e., a more typical muricid-like morphology), but this was still variable depending on local shell thickness at the site of the drillhole. Thus, £. sulcidentata joins the growing list of muricid gastro- pods capable of drilling beveled naticid-like drillholes (see also Edward et. al., 1992; Gordillo and Amuchas- tegui, 1998; Carriker and Yochelson, 1968). Predators selected drilling sites away from the prey shell margins for attacks on bivalves and _ slipper limpets. However, 50% of barnacle prey consumed (n Page 29 = 8) were edge drilling attacks between wall plates. Only one attack on B. eburneus was a drillhole through a wall (lateral plate), and the remainder (n = 3) were drillholes through the beak (scutum). The more refined attack behaviors used against barnacles suggests that E. sulcidentata may specialize on this prey type in the wild. For drilling attacks recorded on the slipper limpets C. depressa and B. aculeatus, 42% (5/12) resulted in incomplete drillholes and unsuccessful attacks. No obvious defensive responses by the slipper limpets were observed during the course of the experiment, although all were mobile and periodically changed their position on the Mercenaria shell or even moved onto the interior of the plastic box. The ratio of incomplete drillholes to total drilling attempts was higher for the spiny Bostrycapulus (67%, 2/3) than the smooth surfaced Crepidula (33%, 3/9), although these numbers are not statistically significant. It is notable, however, that the predator offered the slipper limpets selectively drilled all of the non-spiny C. depressa first. Only when the nine unornamented C. depressa had been eliminated from its box did E. sulcidentata begin to attack and drill the spiny Bostrycapulus. Acknowledgments. We thank Mr. Jospeh Krivanek for assisting us in the field and the USF Geology Deparment for supporting this class project. LITERATURE CITED ABE, N. 1983. Breeding of Thais clavigera (Kuster) and predation of its eggs by Cronia margariticola (Broderip). Pp. 381-392 in B. Morton & D. Dudgeon (eds.), The malacofauna of Hong Kong and southern China III. Hong Kong University Press: Hong Kong. CARRIKER, M. R. 1961. Comparative functional morphology of boring mechanism in gastropods. American Zoologist 1:263—266. CARRIKER, M. R. 1981. Shell penetration and feeding by naticacean and muricacean predatory gastropods: a synthesis. Malacologia 20:403—-422. CARRIKER, M. R. & G. L. GRUBER. 1999. Uniqueness of the gastropod accessory boring organ (ABO): comparative biology, an update. Journal of Shellfish Research 18:579— 595. CARRIKER, M. R., J. G. SCHAADT & V. PETERS. 1974. Analysis by slow-motion picture photography and scanning electron microscopy of radular function in Urosalpinx cinerea follyensis (Muricidae, Gastropoda) during shell penetration. Marine Biology 25:63—76. CARRIKER, M. R. & E. L. YOCHELSON. 1968. Recent gastripod boreholes and Ordovician cylindrical borings. U.S. Geol. Surv. Prof. Pap. 593-B, 26 pp. COLLIN, R. 2003. Phylogenetic relationships among Calyp- traeid gastropods and their implications for the biogeog- raphy of marine speciation. Systematic Biology 52:618— 640. DUNKIN, S. DE B. & R. N. HUGHES. 1984. Behavioural components of prey-selection by dogwhelks, Nucella lapillus (L.) feeding on barnacles, Semibalanus balanoides Page 30 The Velicer Vols 505 Nom (L.) in the laboratory. Journal of Experimental Marine Biology and Ecology 79:91—103. EDWARD, J. K. P., M. X. RAMESH & K. AYAKKANNU. 1992. Comparative study of holes in bivalves, chipped and bored by the muricid gastropods Chicoreus ramosus, Chicoreus virgineus and Murex tribulus. Phuket Marine Biological Center Special Publication 11:106—110. GORDILLO, S. & S. N. AMUCHASTEGUI. 1998. Estrategias de depredacion del gastrOpodo perforador Trophon geversia- nus (Pallas) (Muricoidea:Trophonidae). Malacologia 39: 83-91. IsHIDA, S. 2001. An analysis of feeding aggregations in intertidal muricids: species-specific mode of foraging- initial predation and parasitism. Asian marine biology 18: 1-13. ISHIDA, S. 2004. initial predation and parasitism by muricid whelks demonstrated by the correspondence between drilled holes and their apparent enveloper. Journal of Experimental Marine Biology and Ecology 305:233-245. MATSUKUMA, A. 1977. Notes on Genkaimurex varicosa (Kuroda, 1953) (Prosobranchia: Neogastropoda). Venus 36:81-88. ; Morton, B. 1994. Prey preference and method of attack by Rapana bezoar (Gastropoda: Muricidae) from Hong Kong. Pp. 309-325 in B. Morton (ed.), The malacofauna of Hong Kong and southern China III. Hong Kong University Press: Hong Kong. PERRY, D. M. 1985. Function of shell spine in the predaceous rocky intertidal snail Acanthina spirata (Prosobranchia: Muricacea). Marine Biology 88:51—58. PHILLIPS, B. F. 1969. The population ecology of the whelk Dicathais aegrota in western Australia. Australian Journal of Marine and Freshwater Research 20: 225-265. RADWIN, G. E. & H. W. WELLS. 1968. Comparative radular morphology and feeding habits of muricid gastropods from the Gulf of Mexico. Bulletin of Marine Science 18: 72-85. ROBERTSON, R. 1970. Review of the predators and parasites of stony corals, with special reference to symbiotic prosobranch gastropods. Pacific Science 24:43—54. ROLLER, R. A., J. D. RICKETT & W. B. STICKLE. 1995. The hypobranchial gland of the estuarine snail Stramonita haemastoma canaliculata (Gray) (Prosobranchia: Murici- dae): a light and electron microscope study. American Malacological Bulletin 11:177—190. TAYLOR, J. D. 1976. Habitats, abundance and diet of muricacean gastropods at Aldabra Atoll. Zoological Journal of the Linnean Society 59:155—-193. WARD, J. 1965. The digestive tract and its relation to feeding habits in the stenoglossan prosobranch Coralliophila abbreviata (Lamark) [sic]. Canadian Journal of Zoology 43:447-464. WELLS, H. W. 1958. Feeding habits of Murex fulvescens. Ecology 39:556—558. WEST, D. J., E. B. ANDREws, A. R. MCVEAN, D. J. OSBORNE & M. C. THORNDYKE. 1994. Isolation of serotonin from the accessory salivary glands of the marine snail Nucella lapillus. Toxicon 32:1261—1264. Wu, S. K. 1965a. Comparative functional studies of the digestive system of the muricid gastropods Drupa ricinia and Morula granulata. Malacologia 3:211—233. The Veliger 50(1):31—38 (March 11, 2008) ames een © CMS, Inc., 2007 A Large New Species of Lobatus (Gastropoda: Strombidae) from the Neogene of the Dominican Republic, with Notes on the Genus BERNARD M. LANDAU International Health Centres, Av. Infante D. Henrique 7, 8200 Albufeira, Portugal and Assistant Researcher, Departamento e Centro de Geologia, Universidade de Lisboa, C6. Campo Grande, 1749-016 LISBOA, Portugal (e-mail: bernielandau@sapo.pt) GIJS C. KRONENBERG Milieu Educatie Centrum, P.O. Box 435, NL-5600 AK Eindhoven, The Netherlands (e-mail: gijs.kronenberg@tiscali.nl) GREGORY S. HERBERT Department of Geology, University of South Florida, Tampa, FL, 33620, USA (e-mail: gherbert@cas.usf.edu) Abstract. A very large new stromb is described from the Neogene of the Dominican Republic, Lobatus vokesae sp. nov. Lobatus is used here as a genus to include the Tropical American clade of species previously placed in the subgenus Tricornis. The new species has characters in common with all the various subgenera proposed within the genus, making subgeneric assignment of this early member of the genus Lobatus undesirable. Key Words: Gastropoda, Lobatus, new species, systematics, Neogene, Dominican Republic. INTRODUCTION The Strombidae are a group of tropical to subtropical gastropods, predominantly inhabiting the intertidal and subtidal zones, feeding on macroalgae and epiphytes (Robertson, 1961; Berg, 1975). Species in this family are conspicuous because of their medium- sized to very large, solid, heavy shells. The early and late Miocene deposits on the island of Hispaniola, and more specifically the outcrops occurring along the Cibao valley of the Dominican Republic, are well known for their fauna of Strombi- dae. Sowerby (1850) was the first to describe fossil mollusks from these rich localities and named three species of Strombus. Subsequent workers such as Maury (1917) and Pilsbry & Johnson (1917) have brought the number of strombid taxa known from these deposits to nine (Table 1). However, this is far from being a comprehensive list of the strombid taxa occurring in the Dominican Republic. Our own collections (BL) include probably four undescribed species, the most spectacular of which is described in this paper. SYSTEMATIC DESCRIPTION Genus Lobatus Iredale, 1921 Type species: (by monotypy, Iredale, 1921: 208): - Strombus bituberculatus Lamarck, 1822: 690. Junior subjective synonym of S. raninus Gmelin, 1791: 3511. Recent, West Indies and Florida. Remarks: Throughout the Neogene, the Strombidae have formed an important part of tropical American assemblages, especially in the western Atlantic, where they diversified into numerous species and formed several distinct species groups. Classically, Neogene tropical American Strombus species have been placed in three genera or subgenera; Strombus (s.s.) [type species S. pugilis Linnaeus, 1758, by subsequent designation, Recent, western Atlantic], Lentigo Jousseaume, 1886 [type species Strombus lentiginosus Linnaeus, 1758, by monotypy, Recent, East Africa], and Tricornis Jous- seaume, 1886 [type species Strombus tricornis Lightfoot, 1786, by monotypy, Recent, Indo-Pacific]. The new species described in this paper belongs to the last group. Page 32 The Veliger; Vol: 50) Nomi Table 1 Preliminary list of strombid taxa so far recorded from the Neogene Dominican Republic. Column 1: the original name under which the taxon was described; column 2 the formation in which it occurs (recorded from literature and BL collections); column 3: original references and subsequent references with figures. Sowerby’s (1850) type material was illustrated by Pflug (1961). Taxon Formation Reference Strombus haitensis Sowerby, 1850 Strombus proximus Sowerby, 1850 Strombus bifrons Sowerby, 1850 Gurabo Strombus ambiguus Sowerby, 1850 Unknown Strombus pugiloides Guppy, 1866 ?Cercado Strombus maoensis Maury, 1917 ?Mao Strombus galliformis Pilsbry, 1917 Strombus dominator Pilsbry, 1917 Gurabo Strombus (Lentigo) cf. raninus Gmelin, 1791 Gurabo Gurabo and Cercado Gurabo and Cercado Sowerby, 1850, p. 48, pl. 9, fig. 7 Pflug, 1961, p. 26-27, pl. 3, figs 1, 2, 3, 5, 6, 8 Sowerby, 1850, p. 48, pl. 9, fig. 8 Pflug, 1961, p. 24-26, pl. 2, figs 5, 6, 8, 9, 10 Sowerby, 1850, p. 48, pl. 9, fig. 9 Pflug, 1961, p. 27-28, pl. 4, figs 1-4, 7, 8, 10 Sowerby, 1850, p. 48 Pflug, 1961, p. 28-29, pl. 4, figs 5, 6, 9 Maury, 1917, p. 120, pl. 20, fig. 6 Pflug, 1961, p. 23-24, pl. 3, figs 4, 7 Maury, 1917, p. 120, pl. 21, fig. 1 Pilsbry, 1917, p. 170 Pilsbry & Johnson, 1922, p. 366, pl. 31, figs 1-2 Pilsbry, 1917, p. 170 Pilsbry & Johnson 1922, p. 366, pl. 32, figs 1, 9 Jung & Heintz, 2001, p. 44, fig. 19 The systematics of Tricornis, however, require revision. A molecular phylogeny of strombids shows that Tricornis is polyphyletic and comprised of separate, distantly related tropical American and Indo-Pacific clades (Latiolais et al., 2006). Kronenberg & Lee (2007) argued that Lobatus Iredale, 1921 [type species Strombus bituberculatus Lamarck, 1822, by monotypy, Recent Caribbean (junior subjective syno- nym of S. raninus Gmelin, 1791)] is the first available name for the tropical American group previously known as Tricornis (sensu Abbott, 1960). In this paper, we treat Lobatus as a full genus, as this clade is separated from all other strombids, including Strombus (s.s.), by one of the longest and_ best- supported branches in the strombid molecular tree. Notably, this conflicts with the tree topology of strombids inferred from anatomical data (Simone, 2005). However, the Latiolais et al. (2006) phylogeny is based on an analysis of nearly three times as many strombid taxa and at least twice the number of phylogenetically informative characters as Simone’s (2005) tree, and is less likely to change as more data are added. Whether and how the Lobatus group should be divided into subgenera is not entirely clear with the data at hand. Based on shell features, Petuch (2004) subdivided the tropical American species of Tricornis into several subgenera: Aliger Thiele, 1929 [type species Strombus gallus Linnaeus, 1758, by monotypy, Recent, Caribbe- an], Eustrombus Wenz, 1940 [type species Strombus gigas Linnaeus, 1758, by original designation, Recent, Carib- bean], Macrostrombus Petuch, 2004 [type species Strom- bus costatus Gmelin, 1791, by original designation, Recent, Caribbean] and Titanostrombus Petuch, 2004 [type species Strombus goliath Schroter, 1805, by original designation, Recent, Brazil]. It should be noted that Petuch (1994) regarded all these as subgenera of Strombus, and employed Lobatus in the same fashion, viz. a subgenus of Strombus. The molecular phylogeny of Latiolais et al. (2006) shows S. gallus, S. gigas and S. costatus to be very closely related. The genus Lobatus Iredale, 1921 was introduced without a description. Lobatus is defined here as a clade of medium sized to very large strombs with widely expanding, non digitated, outer lips, and a glazed outer edge of the rim of the outer lip, not bent toward the columella when reaching maturity, usually with strong spiral sculpture on the last whorl. This clade is in Recent times restricted to the Caribbean and Panamic faunal Provinces. It is first recorded from the Lower Miocene Chipola Formation of Florida as Strombus chipolanus Dall, 1890 (Gardner, 1947; Petuch, 2004) and the Middle Miocene Baitoa Forma- tion of the Dominican Republic by another unde- scribed species (Bernard Landau unpublished data). Lobatus is represented in the Late Miocene to Early Pliocene Dominican Republic assemblages by S. haitensis, S. galliformis, S. dominator, S. raninus and Lobatus vokesae sp. nov. Strombus maoensis Maury, 1917 was based on a single incomplete shell, and despite intensive collecting we have found no further specimens. Maury (1917) compared S. maoensis to S. gallus, and it probably represents a species of Lobatus. Strombus ambiguus Sowerby, 1850 is also based on a B. M. Landau et al., 2007 juvenile specimen of a Lobatus species (lectotype illustrated by Pflug, 1961). Abbreviations: The following abbreviations are used: NMB = Naturhistorisches Museum Basel localities; TU = Tulane University localities; NHMW = Naturhistorisches Museum in Wien (Austria) collection number; BL coll. = Bernard Landau collection. SYSTEMATIC DESCRIPTION Genus Lobatus Iredale, 1921 Type species: (by monotypy, Iredale, 1921: 208): - Strombus bituberculatus Lamarck, 1822: 690 QGunior subjective synonym of S. raninus Gmelin, 1791: 3511), Recent, West Indies and Florida. Lobatus vokesae Landau, Kronenberg and Herbert sp. nov. (Figures 1-7) Etymology: We have great pleasure in naming this magnificent shell in honor of Emily Vokes for her enormous contribution to Caribbean Neogene paleontology. Description: (Based on holotype and paratype) Shell very large, solid, when complete reaching at least 270 mm high. Protoconch not known. Seven tele- oconch whorls preserved. Spire broadly conical, weakly coeloconoid in profile, spire whorls depressed in holotype; fifth and penultimate whorls slightly elevated and roundly shouldered in paratype. Sculpture on early teleoconch whorls of small rounded tubercles placed immediately above abapical suture, crossed by numer- ous fine spiral threads. Abapically tubercles become weaker, subobsolete on fourth whorl, and _ spiral threads broaden to form relatively narrow, flattened, subequal spiral cords. Suture impressed, crenulated around tubercles on early teleoconch whorls, weakly undulating abapically. Last whorl greatly inflated, bearing three large, roundly pointed tubercles at shoulder, first tubercle placed opposite (to left of) aperture, third on dorsum, second tubercle midway between other two. Dorsal tubercle very strongly developed, first tubercle slightly smaller, intermediate tubercle weakest. Spiral sculpture of broad, flattened, primary spiral cords, only clearly developed on mid- portion of last whorl, where there are 10-13 cords, and numerous irregular flattened secondary cords. Growth lines prominent on portions of last whorl, interrupting secondary cords, giving a somewhat reticulate aspect to parts of dorsum. Outer lip not thickened, greatly expanded, its adapical end extended above height of apex (when outer lip complete) and expanded medially to join ventral midline. Outer edge of lip and strombid Page 33 notch not preserved. Parietal wall smooth. Base of columella strongly bent backwards. Siphonal canal open, relatively long and broad, bent slightly to right and weakly posteriorly recurved. Holotype: NHMW 2007z0161/0001 Dimensions of holotype: Length 220 mm; dorso-ventral height (restored) 165 mm; width 190 mm (Figures 1— 3). Type locality: Rio Cana, area equivalent to NMB 16832/16833 and TU 1230, Cercado Formation (late Miocene) (Saunders et al., 1986, text-figure 15). Material: Holotype; and | Paratype (B. Landau coll.), length 264 mm; dorso-ventral 150 mm; width (incom- plete outer lip) 180 mm (Figures 4-7); locality Canada de Zamba, off Rio Cana, area equivalent to NMB 16817 and TU 1354, Gurabo Formation (base of the Pliocene) (Saunders et al., 1986, text-figure 15). Remarks: This species is based on two adult specimens from different localities along the Rio Cana, the holotype from beds of late Miocene age, the paratype from basal Pliocene beds. Neither of the specimens is perfect; both are missing the outer part of the outer lip (more complete in the holotype), and the holotype has the top of the dorsal tubercle abraded. The paratype shows signs of damage during life, possibly as a result of attack by a predator, and subsequent repair, with an irregular fracture line running the whole length of the last whorl. There are some differences between the two speci- mens; the paratype when complete would have been the larger shell. It has a slightly more elevated spire than the holotype, and the tubercles at the shoulder of the last whorl are even more massive than in the holotype. Comparisons: Lobatus vokesae sp. nov. is similar in size to the Recent Lobatus gigas, and both species have a broadly expanded but not thickened outer lip. The character of their spires, however, is quite different, as it is much more elevated in L. gigas than in L. vokesae n. sp., with all the spire whorls bearing pointed tubercles and a greater number of more pointed tubercles on the shoulder of the last whorl. The first record of L. gigas is from the Bowden Formation of Jamaica (Jung, 1971). The Bowden Beds are usually considered late Miocene to early Pliocene (Berggren, 1993) or early Pliocene (Bolli & Bermudez, 1965: Bolli & Premoli Silva, 1973; Jung & Heitz, 2001), although Aubry (1993) placed them in the early late Pliocene (calcareous nannoplankton zone NNI16). It has been suggested to us that the Bowden Formation is an olistostrome, which would account for these different ages (Oliver Macsotay, personal communication, The Veliger, Vol. 50, No. 1 Figures 1-3. Lobatus vokesae Landau, Kronenberg and Herbert sp. nov. Holotype, NHMW 2007z0161/0001. Locality: Rio Cana, area equivalent to NMB 16832/16833 and TU 1230, Cercado Formation (late Miocene) (Saunders et al., 1986, text-figure 15). Length 220 mm; dorso-ventral height (restored) 165 mm; width 190 mm. 2007). Lobatus gigas has not been found in the Dominican Republic assemblages. The Recent Lobatus goliath Schréter, 1805 has an even larger shell, also with a non-thickened outer lip, which is even more widely expanded than in the L. vokesae sp. nov. or L. gigas. The spire of L. goliath is imilar to that of our new species; depressed, devoid of ‘tubercles (or almost so) and with a coeloconoid profile, but the tubercles on the last whorl are more numerous and far less strongly developed than in L. vokesae sp. nov. We are not aware of any fossil record for L. goliath. Lobatus williamsi (Olsson & Petit, 1964) from the late Pliocene of Florida, allocated to Titanostrom- bus by Petuch (1994), also lacks the large shoulder tubercules on the last whorl of our new species, but it has some distinct knobs on the shoulder of the B. M. Landau et al., 2007 Page 35 Figures 4-7. Lobatus vokesae Landau, Kronenberg and Herbert sp. nov. Paratype, BL coll. Locality: Canada de Zamba, off Rio Cana, area equivalent to NMB 16817 and TU 1354, Gurabo Formation (base of the Pliocene) (Saunders et al., 1986, text-figure 15). Length 264 mm; dorso-ventral 150 mm; width 180 mm. Page 36 The Veliger, Vol. 50, No. 1 penultimate whorl (Petuch, 1994: pl. 20, fig. A) that are not present in L. vokesae sp. nov. The characters of the tubercles and the depressed spire, almost devoid of sculpture, are similar to those of the Recent Florida species Lobatus costatus Gmelin, 1791. However, in L. costatus the outer lip is not so greatly expanded, especially in the adapical portion, and is very strongly thickened. L. costatus is also a smaller species and lacks the complex spiral sculpture of primary and secondary cords present on the last whorl of L. vokesae. Petuch (1994) described a subspecies Strombus (Macrostrombus) costatus griffini Petuch, 1994 from the Late Pliocene of Florida. The shell illustrated (Petuch, 1994, pl. 19, fig. H) and numerous topotypes in the BL collection all have much weaker tubercles than the Recent shell or L. vokesae and we agree that they are not conspecific. However, several other Strombus (Macrostrombus) species were described in the same publication, very similar to this subspecies, and a review of these taxa is beyond the scope of this work. Lobatus costatus occurs fossil in the Pleistocene Tortuga Formation of Cubagua Island, Venezuela (Bernard Landau unpublished data). Within the Dominican assemblage, Lobatus vokesae sp. nov. is most similar to L. dominator in the character of its very prominent dorsal tubercle. However, L. dominator has a much smaller shell (less than half the size of L. vokesae), the primary spiral cords on the last whorl are more distinct, and the shape of the outer lip is quite different. We (BL coll.) have six specimens of L. dominator from the Cercado and Gurabo Formations of the Dominican Republic. These specimens are highly variable, and in the largest the adapical extremity of the outer lip is developed into an open digitation, similar to but not as elongated as that seen in the Recent Caribbean species L. gallus (Linnaeus, 1758). Strombus dominator delabechei Rutsch, 1931, described from the Bowden Beds of Jamaica, probably falls within the range of variation of Dominican Republic specimens. The shells of Lobatus vokesae sp. nov. demonstrate characters intermediate between those of the subgenera Eustrombus (very large, greatly expanded and not thickened outer lip), Macrostrombus (ribbed last whorl sculpture, large tubercles at the shoulder) and Titanos- trombus (very large size, combined with a_ low, coeloconoid spire) as recognized by Petuch. Therefore, it seems undesirable to assign L. vokesae, an early species of Lobatus, to any subgenus within the genus Lobatus. In the more Recent fossil record fairly well defined lineages can be recognized within Lobatus. The origins of the genus Lobatus are unclear at present, but it is likely that it arose from a species which would be allocated to Persististrombus Kronenberg & Lee, 2007 Within Persististrombus there are some tendencies towards Lobatus-like species (Harzhauser & Kronenberg, in prep.). Geological and environmental setting: The holotype is from the Rio Cana, area equivalent to NMB 16832/ 16833 and TU 1230, Cercado Formation (late Mio- cene) (Saunders et al., 1986, text-figure 15). This is a 50 cm thick bed with closely packed small molluscs in which Astraea Réding, 1758 and Tegula Lesson, 1835 predominate. Other common gastropods are Erosaria Spurca (Linnaeus, 1758), Pachycrommium guppyi (Gabb, 1873), Polinices subclausa (Sowerby, 1850), Neverita (Hypterita) nereidis (Maury, 1917) Semicassis reclusum (Guppy, 1873) and Chicoreus cornurectus (Guppy, 1876). These probably represent shallow inshore conditions. The paratype is from the Canada de Zamba locality, a tributary of the Rio Cana, area equivalent to NMB 16817 and TU 1354, Gurabo Formation (base of the Pliocene) (Saunders et al., 1986, text-figure 15). This locality has a rich and varied gastropod fauna with no particular group predominant. Corals are common and represent a reef structure probably less than 30 m in depth (Saunders et al., 1986). Occurrence: Known only from the late Miocene and early Pliocene Cercado and Gurabo Formations of Rio Cana and its tributary Canada de Zamba, Dominican Republic. DISCUSSION Although the development of a dorsal tubercle is widespread in Recent species of Lobatus, the occur- rence of this feature in L. vokesae is one of the earliest examples within the genus. The function of the dorsal shell protuberance is almost certainly anti-predatory, with a primary role in helping the animal to right its shell after being turned over by predatory fish, crabs, and octopus (Berg, 1975). The large dorsal tubercle forces the overturned shell to lean to either side, which reduces the time and extent to which the animal must extend its soft foot outward and unprotected to right the shell (Savazzi, 1991; see also Carefoot and Donovan, 1995). Selection for a prominent dorsal tubercle should be greatest in larger strombs with a flaring lip, and L. vokesae was one of the largest early strombs. Berg (1975) demonstrated that larger strom- bids are exposed for a longer period of time during righting due, in part, to their own weight and the broad, heavy lip. Interestingly, the presence or absence of the dorsal tubercle varies interspecifically as well as intraspecifi- cally in strombids, e.g., Persististrombus granulatus (Swainson, 1822). An example of the variability of shoulder knobs within a single species of Lobatus is illustrated by the case of L. fetus (Jung & Heitz, 2001), described from the late Pliocene Escudo de Veraguas Formation, Bocas del Toro area of Panama. In our opinion this is based on a specimen of L. raninus in B. M. Landau et al., 2007 which the large dorsal shoulder knob is not developed. This variability can also be observed today, albeit uncommonly, in Recent specimens (Gis Kronenberg, personal observation) and in Pleistocene fossil and Recent L. costatus (Gregory Herbert, personal obser- vation). Therefore, a division of Lobatus into subgenera based solely on this sculptural element is unwarranted. The Plio-Pleistocene radiation of Lobatus species in Florida resulted in a cohort of species that all had weaker tubercles than the fossil L. vokesae and living L. costatus or L. raninus, or had lost them altogether as in the cases of Strombus (Macrostrombus) hertweckorum Petuch, 1991 and Strombus (Macrostrombus) leidyi Heilprin 1886. Whatever changes in predatory patterns led to the loss of tubercles in the Floridian assemblages seem to have affected strombs as a whole, as the genus Strombus (s.s.), which is also greatly diversified in the Plio-Pleistocene of Florida, shows a similar pattern, with a radiation of Pliocene species with no tubercles on the last whorl (see Petuch, 1994; Hargreaves, 1995). The actual number of species within this radiation cannot be commented with certainty at present, as the numerous taxa described and illustrated by Petuch (1994) require revision. REFERENCES ABBoTT, R. T. 1960. The genus Strombus in the Indo-Pacific. Indo-Pacific Mollusca 1:33-146. Aubry, M. P. 1993. Calcareous nannofossil stratigraphy of the Neogene formations of eastern Jamaica. In: E. R. Robinson, J. B. Saunders & R. M. Wright (eds.), Biostratigraphy of Jamaica. Geological Society of Amer- ica Memoir 182: Pp. 131-178. BERG, C. J., JR. 1975. 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The Veliger 50(1):39-47 (March 11, 2008) THE VELIGER © CMS, Inc., 2007 First Record of the Northeastern Pacific Patellogastropod Genus Acmaea from the Miocene of Japan and Its Paleobiogeographic Implications YUKITO KURIHARA AND TOMOKI KASE Department of Geology, the National Museum of Nature and Science (formerly the National Science Museum, Tokyo), 3-23-1 Hyakunincho, Shinjuku-ku, Tokyo, 169-0073, Japan (e-mail: kurihara@kahaku.go.jp; kase@kahaku.go.jp) Abstract. Acmaea mitra Rathke, 1833 is the sole species of the genus Acmaea and inhabits lower intertidal to subtidal rocky shores in the eastern side of the North Pacific Ocean from California to the Aleutian Islands. The discovery of Acmaea mitra from the Upper Miocene of central Japan represents the first fossil record of the species and genus in the western side of the North Pacific and the oldest record of the genus Acmaea. The Late Miocene specimens are definitely referred to Acmaea mitra on the basis of overall shell morphology and shell microstructure. The fossil records strongly suggest that A. mitra became regionally extinct in the western side of the North Pacific by the end of the Late Miocene. A. mitra represents a good additional example of northeastern Pacific restriction, an uncommon biogeographic distribution pattern of marine organisms in the middle-latitude North Pacific Ocean during the late Cenozoic. The causes of the regional extinction of A. mitra in the western side of the North Pacific remain uncertain. INTRODUCTION Both the western side of the North Pacific (WSNP) and the eastern side of the North Pacific (ESNP) have well- documented fossil records of Cenozoic marine mol- lusks that provide basic data for examining the historical development of marine biogeography in this bioprovince. An interesting distribution pattern seen in fossil and modern mollusks in this bioprovince is geographic restriction: taxa living on both sides of the North Pacific during the Neogene subsequently became restricted either to the WSNP (the northwestern Pacific restriction) or’ the ESNP (the northeastern Pacific restriction). Vermeij (1989) surveyed historical biogeo- graphic patterns of cool-temperate mollusks during the Neogene and recognized 15 taxa that reflected the northwestern Pacific restriction. On the other hand, he recognized only a possible case of the northeastern Pacific restriction. Later, Amano (1998) and Kurihara (2007) added several taxa reflecting this pattern. In these studies, the causes of regional extinction in the WSNP have not been examined rigorously. In this study, we present another case of northeastern Pacific restriction recognized in the patellogastropod limpet genus Acmaea. The genus Acmaea accommodates only the single species Acmaea mitra Rathke, 1833. Many species once allocated to Acmaea in the North Pacific have now been referred to other genera of Lottiidae (e.g., Lindberg, 1981, 1986; Sasaki, 1999), and those fossil species described as Acmaea were based solely on shell morphology, a highly convergent character among patellogastropods, so that they need further study for reliable generic allocation. A. mitra inhabits hard substrates from the low intertidal to a depth of 30 m along the ESNP and the eastern Aleutian Islands (Lindberg, 1981). The discovery of A. mitra from the Upper Miocene of Japan reported in this paper is twofold: (1) the fossil record of the genus Acmaea extends back to the Late Miocene, and (2) it shows regional extinction of A. mitra in the WSNP by the end of the Late Miocene. The following institutional abbreviations are used: GMNH (Gunma Museum of Natural History, To- mioka, Gunma, Japan), NSM (National Museum of Nature and Science, Tokyo; formerly National Science Museum, Tokyo, Japan), SDSNH (San Diego Natural History Museum, California, U.S.A.) and UMUT (University Museum, the University of Tokyo, Japan). STRATIGRAPHY AND ASSOCIATED FAUNA The three Acmaea specimens described in this paper are found in the molluscan fossil collection of GMNH (PI2258—2260). They were collected at locality HNO8& [= locality C of Kato (2001)], a right bank of the Usui River, Minakuchi, Annaka City, Gunma Prefecture Page 40 (36°19'11"N, 138°53'12”E) in a fossiliferous pebbly medium- to coarse-grained sandstone bed in the lower part of the Itahana Formation. The Itahana Formation is the uppermost regressive unit of the Miocene Annaka Group (Takahashi & Hayashi, 2004), and is stratigraphically divided into the lower marine and upper non-marine units (Oishi & Takahashi, 1990). Based on the radiometric and biochronologic analyses of the underlying Haraichi Formation, Takahashi & Hayashi (2004) estimated the Itahana Formation as early Late Miocene age (ca. 11.0 Ma). The lower unit of the Itahana Formation contains well-preserved marine mollusks of about 80 species (Kurihara, 2000). This assemblage is typical of Shiobara-type molluscan fauna (Iwasaki, 1970; Chin- zel, 1978), Late Miocene temperate shallow-water associations characteristic in central and northern Honshu, Japan (Chinzei, 1986). Mollusks associated with Acmaea occupied various shallow marine habitats and include species of the gastropods Charonia, Chlorostoma and Kelletia indicative of temperate, subtidal rocky shores. SYSTEMATIC PALEONTOLOGY Order PATELLOGASTROPODA Lindberg, 1986 Superfamily PATELLOIDEA Rafinesque, 1815 Family LOTTIITDAE Gray, 1840 Genus Acmaea Eschscholtz in Rathke, 1833 Type species: Acmaea mitra Rathke, 1833, subsequent designation by Dall, 1871. Discussion: Lindberg (1986) revised familial and generic level classification of species traditionally assigned to ““Acmaeidae.”” Major changes in his classification are subdivision of ‘“‘Acmaeidae” into Acmaeidae and Lottiudae and the restricted usage of Acmaeidae to a small monophyletic group that includes the monotypic, shallow-water genus Acmaea and the deep-water genus Pectinodonta. The members of this group share three pairs of uniform lateral teeth arranged in a posteriorly diverging inverted V-shape, identical ventral plate morphology, an absence of marginal teeth, similar gross anatomy and the same shell structure belonging to MacClintock’s (1967) shell structure group 15 (Lindberg, 1986). However, Nakano & Ozawa (2004) recently demonstrated that A. mitra and Niveotectura pallida (Gould, 1859) constitute a monophyletic group on the basis of molecular data and similarity of radula, and also that Pectinodonta is clearly unrelated to A. mitra. Nakano & Ozawa (2007) classified A. mitra and \. pallida as a clade within Lottiidae and regarded ithe Veliger, Voly 50. Now Figure 1. 1, 2, Acmaea mitra Rathke from the Upper Miocene Itahana Formation, apical and lateral views, GMNH P12258, locality HNO08, length, 21.0 mm, width, 19.0 mm, height, 15.3 mm. Acmaeidae as a junior synonym of Lottiidae. The validity of this new systematic change needs to be confirmed by rigorous anatomical study because 4A. mitra and N. pallida belong to different shell structure groups and the anatomy of A. mitra has never been studied (Fuchigami & Sasaki, 2005). In this paper, we follow the classification proposed by Nakano & Ozawa (2007) for the familial assignment of Acmaea. Acmaea mitra Rathke, 1833 (Figures 1, 2) Acmaea mitra RATHKE IN ESCHSCHOLTZ, 1833, p. 18, pl. 23, fig. 4; ABBOTT, 1974, p. 29, fig. 145; LINDBERG, 1981, p. 63, fig. 64; LINDBERG & MARINCOVICH, 1986, fig. 2h; LINDBERG, 1988a, fig. 6d. Description: Shell up to 21.0 mm in length, high conical, moderately thick, cap-shaped, with height about 3/4 of major apertural diameter. Apex in anterior 2/5, not curved anteriorly. Aperture subcircular, with length/width ratio 1.23. Anterior slope very weakly convex and other slopes almost straight. Surface devoid of any sculpture except for some concentric, knobby bulges indicative of growth halts, and concentric Y. Kurihara & T. Kase, 2007 Page 41 Figure 2. SEM micrographs of the shell walls in Acmaea mitra Rathke from the Itahana Formation, GMNH PI2259, locality HN0O8, etched by 1% HCl for 30 sec. 1, fractured section cut almost radially from the apex to shell margin showing the outermost complex prismatic layer (M+3), followed by the foliate layer (M+2), outer concentric crossed-lamellar layer (M+1), myostracum layer (M) and inner radial crossed-lamellar layer (M—1); 2, polished surface cut almost commarginally showing the foliate layer (M+2), followed by the outer concentric crossed-lamellar layer (M+1) and myostracum layer (M). growth lamellae. Shell consists of five layers; outermost layer (M+3) of complex prismatic structure, followed by foliate layer (M+2), concentric crossed-lamellar layer (M+1), myostracum (M), radial crossed-lamellar layer (M-1). - Discussion: The suprageneric classification of living patellogastropods is primarily based on radular, gill and other anatomical characters, not preserved in fossil shells. However, shell morphology frequently exhibits convergence and parallelism, which makes the generic classification difficult. Analysis of shell microstructures is a powerful method for classification of fossil species (e.g., Lindberg & Hickman, 1986; Lindberg, 1988a, b; Lindberg & Marincovich, 1988; Lindberg & Squires, 1990; Kase, 1994; Kase & Shigeta, 1996; Lindberg & Hedegaard, 1996). Patellogastropod shells consist of four to six successive layers including the myostracum, and each layer is composed of one of four basic microstructures (prismatic, foliated, crossed, and com- plex crossed) and also of either a microstructure different from adjacent layers, or, where the structure is the same, the two layer’s major structural elements are oriented perpendicular to each other (MacClintock, 1967). This study demonstrated a general consistency between the classification based on soft anatomy and shell structure (MacClintock, 1967), who recognized 17 shell structure groups within the Patellogastropoda. Recently, Fuchigami & Sasaki (2005) added some deep- sea taxa recently accessible by submersible vessels and recognized 20 shell structure groups. They further emphasized the general, but not complete, consistency between the soft anatomy and shell structure. Among the 20 shell structure groups recognized by Fuchigami & Sasaki (2005), the specimens described here belong to their shell structure group P, diagnosed by an outermost irregular spherulitic prismatic layer, fol- lowed by a concentric regular foliated layer, concentri- cally arranged crossed-lamellar layer, the myostracum, and an inner radially arranged crossed-lamellar layer, clearly demonstrating allocation to the genus Acmaea. The Late Miocene specimens from the Itahana Formation do no exhibit any difference in overall shell morphology from those of the modern specimens of Acmaea mitra. The periodic concentric and knobby bulges are seen both in the fossil and modern specimens. The largest specimen from the Itahana is slightly smaller than the common adult size of the modern specimens, but it appears not to be an important distinguishing character. Acmaea sookensis Clark & Arnold (1923) from the Upper Oligocene Sooke Formation of Vancouver Island, British Columbia, Canada is the only fossil form that is sculptured only by concentric and periodic increments similar to those of the present species. This species, however, was reassigned to the genus Patelloida by Lindberg & Marincovich (1988) and therefore their resemblance is only superficial. Distribution: A. mitra inhabits hard substrates of the low intertidal to a depth of 30 m along the Pacific side of North America from the warm-temperate sea of Isla San Martin, Baja California, Mexico (30°30’N) in the south to the cool-temperate sea of Umnak Island, eastern Aleutian Islands, Alaska (53°N) to the north (Lindberg, 1981; Vermeij et al., 1990). Recently, Golikov et al. (2001) recorded this species from the Sea of Okhotsk for the first time, but this record is Page 42 the Velicer, Vol s05 Nomi Table 1 Comparisons of shell and radular characters, habitat and feeding between Acmaea mitra and Niveotectura pallida. Acmaea mitra Niveotectura pallida Shell Color White! White? Maximum diameter ca. 30 mm! ca. 60 mm? Profile High! High? Subcentral! Subcentral’ Structure group Group P? Group C* Sculpture Concentric growth lines! Radial ribs and concentric growth lines? Radula formula 0-3-0-3-0! 0-3-0-3-0* Habitat Substratum Hard bottom! Hard bottom? Bathymetric range Lower intertidal to 30 m! Lower intertidal to 70 m° Geographic range Aleutians to Baja California! Sakhalin and Kuriles to central Japan, Korea, Maritime Territory” Food Coralline algae® Coralline algae’ References: 'Lindberg (1981); *Sasaki (2000); *Fuchigami & Sasaki (2005); *Sasaki (1998); °Sasaki (2006); “Padilla (1985); "Fujita (1992). . based on the misidentification of Erginus moskalevi (Golikov and Kussakin, 1972) (B. Sirenko, pers. comm.). Therefore, the geographic distribution of this species is currently restricted to the eastern North Pacific. PALEOBIOGEOGRAPHIC IMPLICATIONS Acmaea mitra is a subtidal species widely distributed along the ESNP from California to the eastern Aleutian Islands (Lindberg, 1981). In contrast, the fossil record of the genus Acmaea is represented only by A. mitra and was considered restricted to the ESNP. 4. mitra occurs in the Pleistocene deposits of California (Grant & Gale, 1931; Valentine, 1961; Marincovich, 1976), and its oldest form is a well-preserved specimen (SDSNH 24351) from the lower part of the San Diego Formation of California, which is dated as middle Pliocene (ca. 3.5 Ma; T. A. Deméré, pers. comm.). Putative oldest forms of Acmaea in the ESNP are Acmaea clarki Van Winkle (1918) from the Oligocene of Washington and Acmaea? cf. A. mitra in the faunal list of the Upper Miocene Towsley Formation, the Ventura Basin of California (Winterer & Durham, 1962), but their identification cannot be confirmed because both species have never been studied rigorous- ly. Therefore, the Late Miocene specimens in Japan represent the oldest fossil record and the first occur- rence in the WSNP of this unique genus and species, and the most parsimonious view is that A. mitra originated in the WSNP during the Late Miocene and later migrated to the ESNP. On the other hand, there is no reported occurrence of A. mitra in the Phocene and Pleistocene of Japan, in pite of the presence of many fossiliferous localities iclding patellogastropod limpets as stated below. This strongly suggests that A. mitra in the WSNP might have become extinct by the end of the Miocene. Vermeij (1989) discussed the origins of various biogeographic patterns seen today among late Neogene cool-water marine mollusks in the North Pacific and North Atlantic. The distribution pattern of A. mitra can be categorized into his “northeastern Pacific restriction,’ where species previously with an amphi- Pacific distribution have become restricted to the ESNP. Molluscan taxa showing this distribution pattern include the lucinid bivalve Epilucina californica (Conrad, 1837), the venerid bivalves Humilaria Grant & Gale, 1931 and Compsomyax Stewart, 1930, the myid bivalve Platyodon Conrad, 1837, and the muricid gastropod Nucella shiwa (Chinzei, 1961) (Amano, 1998; Kurihara, 2007). As far as we are aware, the following genera can be added to the northeastern Pacific restriction taxa: the turrid gastropod Mega- surcula Casey, 1904 and the cymatiid gastropod Mediargo Terry, 1968. Megasurcula is still extant in the ESNP, but in the WSNP this genus became extinct by the end of the Late Miocene (Oyama, 1954). Mediargo became extinct by the Pliocene in the ESNP, whereas it persisted only until the end of the Late Miocene in the WSNP (Smith, 1970). From _ the biostratigraphic point of view, the majority of cool- and mild-temperate marine molluscan clades in Japan range from the Late Miocene to Pliocene, suggesting that only a few clades became extinct during the Late Miocene. Aside from Acmaea, Megasurcula and Mediargo, the only molluscan groups extinct during the Late Miocene are the pectinid genera Nanaochla- mys Hatai & Masuda, 1953 and Mivagipecten Masuda, 1952 (Masuda, 1986; Matsubara, 1996). Patterns of geographical restriction provide a clue for understanding the causes of extinction. If a species Y. Kurihara & T. Kase, 2007 Page 43 persisted in one area while it disappeared in another, the possible causes of extinction can be attributed to the factors or events by which the two areas differ (Vermeij, 1989). Similarly, if a species disappeared in one area while its close relative persisted in the same area, the possible causes of extinction can be attributed to the factors by which the two species differ. A. mitra, now restricted to the ESNP, occupies the same habitat with, and is the closest relative to N. pallida, which is restricted to the WSNP (Nakano & Ozawa, 2004). Although these two species belong to different shell structure groups and have different external sculpture, they are almost identical in shell form and color, radular morphology, and feeding strategy (Table 1). Therefore, both A. mitra and N. pallida are regarded as ecological counterparts and may have responded similarly to changing environments in the geological past. We undertook an extensive survey of the geographic and stratigraphic distributions of N. pallida in Japan (Figure 3). In this survey we treated Niveotectura shigaramiensis (Makiyama, 1927) as a junior synonym of N. pallida. Miocene specimens assignable to N. pallida are from the Ginzan Formation of Yamagata Prefecture (Nomura & Zinbo, 1937; NSM PM18322) and the Koshitomaezawa Formation of Iwate Prefec- ture (NSM PM17596), both in northeast Honshu. The Ginzan Formation has been dated as the late Middle Miocene by planktonic foraminifera and diatom biostratigraphy (Sato, 1986), and the Koshitomaezawa Formation as the Middle Miocene or the early Late Miocene based upon radiometric dating of its overlying unit (Suto & Ishii, 1987). The shell of the Ginzan specimen consists of a thick, outer complex prismatic layer and a thin, inner concentric crossed-lamellar layer that belongs to Fuchigami & Sasaki’s (2005) Group P as does the modern N. pallida. The specimen described by Yokoyama (1925b) from Sakae in Nagano Prefec- ture, which Marincovich & Lindberg (1988) regarded as from the Upper Miocene Ogawa Formation, is now believed to be from the Lower Pliocene Shigarami Formation (e.g., Amano &Koike, 1993). Lindberg & Marincovich (1988) noted that the oldest example of WN. pallida was from the Joban coalfield of northeast Honshu, Japan. Yokoyama (1925a) recorded N. pallida from three localities in the Joban coalfield, of which Yunami (Tozenji) is known as a classic (now destroyed) fossil locality of the lower Middle Miocene Kokozura Formation of the Takaku Group (Yanagisawa, 1996). If the occurrence of N. pallida from Yunami is correct, it represents the oldest form of this species. However, we cannot confirm Yokoyama’s (1925a) record because the specimen of N. pallida from Yunami was not illustrated by him and has not been found in the UMUT collection. Therefore, we excluded the occur- rence from Yunami in this discussion. The Pliocene and Hokkaido Pacific Ocean Koshitomaezawa Fm. Ginzan Fm. Itahana Fm. LEGEND Acmaea mitra at, . v9 Miocene Niveotectura pallida © Plio-Pleistocene @ Miocene Figure 3. Geographic and stratigraphic distribution of fossil Acmaea mitra and Niveotectura pallida in Japan. Sources of fossil records of Niveotectura pallida are as follows: 1, Otanoshike Fm. (Akamatsu, 1988); 2, Otoebetsugawa Fm. (Akamatsu, 1987); 3, Setana Fm. (Suzuki, 2002, 2003); 4, Tomikawa Fm. (Sakagami et al., 1966); 5, Narusawa Fim. (Iwai, 1960); 6, Noheji Fm. (Iwai and Siobara, 1969); 7, Sasaoka Fm. (Nomura and Hatai, 1938); 8, Shibikawa and Katanishi Fms. (Ogasawara et al., 1986); 9, Sawane Fm. (Yokoyama, 1926; Omori, 1977); 10, Tanihama Fm. (Amano et al., 1987); 11, Shigarami Fm. (Yokoyama, 1925b; Ma- kiyama, 1927; Nagamori, 1998); 12, Mita Fm. (Fuji and Shimizu, 1991); 13, Zukawa Fm. (Fujii and Shimizu, 1992); 14, Omma Fm. (Yokoyama, 1927; Kaseno and Matsuura, 1965; Matsuura, 1985); 15, Tomioka Fm. [= Dainenji Fm] (Nemoto and O’Hara, 2005); 16, Taga Group [= Dainenji Fm.] (Nemoto and O’Hara, 1979); 17, Hitachi Fm. (Yo- koyama, 1925a; Noda et al., 1995); 18, Shimosa Group (Yokoyama, 1922; O’Hara, 1982); 19, Kazusa Group (Yo- koyama, 1920; Shikama and Masujima, 1969; Baba, 1990); 20, Ninomiya Group (Okumura, 1980); 21, Toyofusa Fm. (Baba, 1990). Records from the Kazusa and Shimosa Groups are too numerous, sO some representative works are cited. Page 44 The Veliger; Vol. 50; Noi Table 2 Ranges of monthly mean sea surface temperature (SST) near the southern and northern distributuion limits of Acmaea mitra and Niveotectura pallida in the North Pacific. Locations Monthly mean SST ranges (°C) Remarks Jueau, Alaska 2.2-10.6' Northern limit of A. mitra Newport Beach, California 14.1-21.1! Southern limit of A. mitra Akkeshi, Hokkaido 0.7-19.1° Northern limit of N. pallida Isozaki, Ibaraki 9.1-21.2? Southern limit of N. pallida References: 'U.S. National Oceanographic Center (2006); "Japan Oceanographic Data Center (2007). Pleistocene occurrences of N. pallida, in contrast, are abundant and distributed widely in central and northern Japan as shown in Figure 3. The fossil record mentioned above indicates that both A. mitra and N. pallida lived in the WSNP during the Late Miocene, and that A. mitra, became extinct there by the end of the Late Miocene whereas N. pallida persists today. Vermeij (1989) hypothesized five major causes that governed the extinction for Neogene marine invertebrates in the North Atlantic and North Pacific: (1) anoxia; (2) regression and habitat loss; (3) reduction in primary productivity; (4) competition and predation; and (5) cooling and warming. The first and second hypotheses are very unlikely because A. mitra inhabited upper subtidal rocky shores where anoxia and habitat loss may hardly have occurred. The third hypothesis has been recognized as a cause of extinction for many Neogene mollusks in the western tropical Atlantic and the eastern temperate North Pacific (see Vermeij, 2001 for review). Vermeij (1989) found that large suspen- sion-feeding bivalves in the ESNP became extinct more than those in the WSNP during the Pliocene, and suggested that reduction or interruption of primary productivity was a possible cause of this extinction in the ESNP. This hypothesis evidently contradicts the distribution pattern of A. mitra. The fourth hypothe- sis—predation and competition as agents of extinc- tion—is well known to be possible causes of extinction for terrestrial organisms but no convincing example has been proposed for marine organisms (Vermeij, 1987, 1989, 2004). The last hypothesis, especially cooling, seems to be the most plausible for the regional extinction of amphi- Pacific marine biota in the WSNP, because the cool- temperate WSNP shows wider annual temperature fluctuations than the ESNP with extensive develop- ment of winter ice (Vermeij, 1978, 1989). However, this hypothesis is unlikely for the selective extinction of A. mitra from the WSNP because this species has wide temperature tolerances the same as WN. pallida in modern seas. The available monthly mean sea surface temperature near the northern- and southernmost distribution areas of A. mitra and N. pallida are almost the same (A. mitra, 2.1-21.1°C; N. pallida, 0.7—21.2°C; Table 2). If A. mitra in the WSNP had become extinct by cooling or warming, N. pallida would have also become extinct at the same time. In the WSNP, cooling and warming events occurred at the latest Miocene (ca. 6—5 Ma) and the Early Pliocene (ca. 54 Ma), respec- tively (e.g., Ogasawara, 1994; Suzuki & Akamatsu, 1994), but N. pallida survived even under these paleoclimatic conditions. G. J. Vermeij (pers. comm.) suggested that cooling is still a possible cause of the regional extinction of A. mitra from the WSNP if this species had a limited geographic range during the Late Miocene in the WSNP: such a small population(s) might have been affected by the environmental deterioration more severely than widely distributed species. However, the poor fossil record of A. mitra and N. pallida during the Late Miocene does not allow us to justify this possibility. In summary, any major hypotheses previously proposed cannot interpret explicitly the development of the unique distribution pattern shown here in A. mitra. However, documentation and accumulation of such examples may contribute toward further under- standing of the origin of various distribution patterns of marine organisms in the North Pacific. Acknowledgments. We thank H. Nakajima (Annaka City) for donation of the fossil specimens of A. mitra to GMNH, Y. Takakuwa (GMNH) for the loan of the specimens in his care, T. A. 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Journal of the Faculty of Science, Imperial University of Tokyo, Section 2, 1:1—23, pls. 1-7. YOKOYAMA, M. 1926. Fossil shells from Sado. Journal of the Faculty of Science, Imperial University of Tokyo, Section 2, 1:249-312, pls. 32-37. YOKOYAMA, M. 1927. Fossil Mollusca from Kaga. Journal of the Faculty of Science, Imperial University of Tokyo, Section 2, 2:165—182, pls. 47-49. The Veliger 50(1):48—S0 (March 11, 2008) THE VELIGER © CMS, Inc., 2007 A New Phaenomenella Fraussen & Hadorn, 2006 (Gastropoda: Buccinidae), from the Andaman Sea KOEN FRAUSSEN Leuvensestraat 25, B-3200 Aarschot, Belgium (e-mail: koen.fraussen@skynet.be) Abstract. A peculiar species from the Andaman Sea is described as Phaenomenella mokenorum sp. nov. The generic placement is based on conchological characteristics of the apical whorls, sculpture and columella. The new species differs from it congeners Ph. inflata (Shikama, 1971), Ph. insulapratasensis (Okutani & Lan, 1994) and Ph. angusta Fraussen & Hadorn, 2006 by having smoother sculpture and a distinct shape. Key Words: Andaman Sea, Gastropoda, Buccinidae, Phaenomenella, new taxon INTRODUCTION The genus Phaenomenella Fraussen & Hadorn, 2006 is characterized by the angular shape of the apical whorls. Two species, Phaenomenella inflata (Shikama, 1971) and Phaenomenella angusta Fraussen & Hadorn, 2006, are known from the East China Sea and off Taiwan. One species, Phaenomenella insulapratasensis (Okutani & Lan, 1994), is known from the South China Sea. The new species here described is from the Andaman Sea, hereby extending the range of the genus into the eastern Indian Ocean. The material reported on in the present study originates from the International Indian Ocean Expe- dition “ANTON BRUUN ’ conducted in 1963. Abbreviations: ANSP Academy of Natural Science, Philadel- phia, U.S.A. KF collection Koen Fraussen, Aarschot, Bel- gium MNHN Muséum national d’Histoire naturelle, Paris, France SYSTEMATICS BUCCINIDAE Rafinesque, 1815 Genus Phaenomenella Fraussen & Hadorn, 2006. Type species: Manaria inflata Shikama, 1971 (type locality: Taiwan) by original designation. Phaenomenella mokenorum sp. nov. Figs. 1-6 fype material: Holotype (55.6 mm) ANSP 291386, paratype | (44.0 mm) and paratype 2 (35.4 mm) ANSP 416212 (from the type locality). Paratype 3 (42.3 mm) and paratype 4 (40.2 mm) (Andaman Sea, southern Myanmar (Burma), west off Twin Island, ANTON BRUUN st. 22B, 10°39’N, 97°06'E, 274-293 m deep, on sand-mud, by dredge, 24/ 3/1963), ANSP 291928. All shells empty or inhabited by hermit crab. Type locality; Andaman Sea, Thailand, E-SE off Phuket Island, ANTON BRUUN stn 17, 07°40'N, 97°08'E, 512-503 m deep, on green-brown clay, by shrimp trawl, 21/3/1963. Range and habitat: Only known from the type material. Andaman Sea, off Thailand and off southern Myan- mar. Description: Shell large for the genus (up to 55.6 mm), thin but solid, color white to pale brownish, shape slender with moderately high spire, body whorl! rather inflated with short siphonal canal. Protoconch rather bulbous, 2.7 mm in diameter, damaged (holotype) or decollate (paratypes), the 2 remaining protoconch whorls are concave and smooth, last whorl slightly angulate below periphery. Transition to teleoconch indistinct. Teleoconch whorls up to 7 in number, laterally flattened. Suture deep, abapically slightly truncate. First 1 1/2 whorls eroded, traces of broad but low and smooth spiral cords occasionally visible, with a fine line as interspace. All following whorls with 10 spiral cords, flattened with shallow, narrow interspaces at second whorl. Spiral cords gradually becoming broad- er, lower and flattened towards penultimate whorl. Body whorl rather smooth, indistinct spiral cords. First 1 1/2 whorls eroded, traces of axial ribs occasionally appearing, last half of second whorl with 8 axial ribs, K. Fraussen, 2007 Page 49 Figures 1-10. 1-6. Phaenomenella mokenorum sp. nov. 1-4. 55.6 mm, holotype, Andaman Sea, Thailand, E-SE off Phuket Island, ANTON BRUUN stn 17, 07°40'N, 97°08’E, 512-503 m deep, ANSP 291386; 5—6. 44.0 mm, paratype 1, same locality, ANSP 416212. 7-8. Phaenomenella inflata (Shikama, 1971) 7. 30.5 mm, Taiwan, TAIWAN 2000, stn CP27, 22°13.3’N, 120°23.5'E, 326 m, MNHN; 8. 33.8 mm, off Suao, Taiwan, dredged, 190 m, KF nr. 0524. 9-10. Phaenomenella insulapratasensis (Okutani & Lan, 1994), 39.2 mm, off Vietnam, trawled by Taiwanese fisherman, KF nr. 1495. Page 50 weak near both sutures, more prominent near periph- ery. Third whorl suddenly smooth, no traces of axial sculpture on subsequent whorls. Aperture semi-oval. Columella strongly twisted, occasionally curved (paratypes), callus thin, smooth. Outer lip thin or slightly thickened, occasionally slightly curled outwards (paratype 1). Outer lip smooth (paratype 1) or with weak traces of internal lirae (holotype), occasionally with 17 or 19 internal lirae (paratypes 3-4). Siphonal canal short, broad, open. Periostracum greenish brown, well adherent, velvety with numerous fine, rather straight (only weakly curved) incremental lamellae running from suture to suture when fresh (holotype), smooth and glossy when eroded (paratype 1). Animal and radula unknown. Remarks: Phaenomenella mokenorum sp. nov. is char- acterized by the elegant shape (slender spire in combination with convex body whorl) and the spiral sculpture of equal strength elements. The generic placement is based on morphological similarities of the distinctive shape of the apical whorls (strongly angular), sculpture and columellar shape, which are all typical for the genus. Phaenomenella inflata from off Taiwan differs by having more convex whorls and a shorter spire in combination with a longer siphonal canal. Phaenomenella insulapratasensis from off Vietnam The Veliger, Vol. 50, No. 1 differs by having a more ovoid shape, a shorter spire, a more solid shell and a smaller adult size. Phaenomenella angusta from off Taiwan and south- ern Japan differs by having a more slender shape with a higher spire and a sharper spiral sculpture. Etymology: Phaenomenella mokenorum sp. nov. is named after the moken, sea-nomads, the indigenous people of the region. Acknowledgments. I am grateful to Paul Callomon (Academy of Natural Science Philadelphia, U.S.A.) for making the type material available for study, to Philippe Bouchet and Virginie Héros (Muséum national d’Histoire naturelle, France) for the loan of material and help, Roland Houart (Belgium) for comments and corrections, Kevin Monsecour for digital images and to David Monsecour (Belgium) for correcting the English text. REFERENCES FRAUSSEN, K. & R. HADORN. 2006. Phaenomenella, a new genus of deep-water buccinid (Gastropoda: buccinidae) with description of a new species from Taiwan. Novapex 7(4):103-109. OKUTANI, T. & T. C. LAN. 1994. A new buccinid whelk collected from Pratas Islets, South China Sea. Bulletin of the Malacological Society of China 18:14. SHIKAMA, T. 1971. On some noteworthy marine Gastropoda from southwestern Japan (III). Science Reports of the Yokohama National University. (section 2) 18:27-35. THE VELIGER i The Veliger 50(1):51-56 (March 11, 2008) © CMS, Inc., 2007 Development of Tylodina fungina Gabb, 1865 (Gastropoda: Notaspidea) from the Pacific Coast of Panama RACHEL COLLIN Smithsonian Tropical Research Institute, Box 0843-03092, Balboa, Ancon, Republic of Panama Abstract. The biology of notaspidean gastropods is not well studied and the development of tylodinoids is almost entirely unknown. Here I report observations on the reproduction and development of Tylodina fungina (Gabb, 1865) from the Perlas Islands on the Pacific coast of Panama. This species lives, feeds, and lays flat egg ribbons on the verongid sponge Suberea azteca (Gomez and Bakus, 1992). The egg ribbons contain hundreds of rows of 80 um eggs, each singly encapsulated in a round 125 um capsule. The ribbon also includes strings of extra-capsular material which is unevenly distributed through the mass. The eggs have equal cleavage and the ciliated “‘trochophore”’ stage is followed by an encapsulated veliger, which has a large, dark-red pigmented mantle organ. At hatching the transparent, left-handed larval shell is 123 um long, and each semicircular velar lobe is unpigmented. There is a distinct operculum, but the eyes and tentacles have not developed. After 3 weeks in culture the larvae had reached a shell length of 162 um and still had no eyes or tentacles. The larvae did not survive to settlement. Key Words: Tylodinidae, Notaspidea, extra-capsular yolk INTRODUCTION The opisthobranch superfamily, Tylodinoidea, con- tains notaspideans with an external, limpet-like shell. The superfamily consists of 2 families, the Umbracu- lidae and the Tylodinidae. Umbraculidae is monotypic with a single species with a worldwide distribution, and the Tylodinidae is comprised of two genera; 7/odina with 5 species and Anidolyta with 2 or possibly 3 species (Willan, 1987). Tylodina is considered “primitive” in the notaspideans. However the monophyly of the notaspideans is contentious, with some families possi- bly including the sister-group of the nudibranchs (Wagele and Willan, 2000). Therefore data on any notaspideans could be useful in testing their monophy- ly as well as -helping to reconstruct character state evolution within the opisthobranchs. Although representatives of both families of tylodi- noids have been examined morphologically, the biology of most of the species remains largely unstudied and the life history is not known for any species of Tylodina (Willan, 1998; Gibbson, 2003). One species each of the five described species of Ty/lodina occurs in Australia, South Africa, the Mediterranean, western Atlantic, and Tropical East Pacific. This unusual biogeographic pattern and the fact that the species are diagnosed with few subtle anatomical features has lead to suggestions that all species of Tylodina should be synonomized under the name T. perversa (Thompson, For correspondence: STRI, Unit 0948, APO AA 34002, USA. e-mail: collinr@si.edu 1970). While disagreeing with this extreme view, Willan (1998) suggests that developmental data could be useful in further demonstrating the validity of the Tylodina species. The only published developmental information for the superfamily is limited to the following observations of T. corticalis and Umbraculum sinicum, each reported for a single spawn of a single female by Thompson (1970) and with some additional information from one other individual of U. sinicum from Ostergaard (1950). Tylodina corticalis is reported to have a bright yellow spiral egg ribbon that is attached flat to the substrate and to contain eggs 98 um in diameter. Umbraculum sinicum deposited a coiled ribbon that contained egg capsules 480-500 um in diameter. Each capsule con- tained 30-45 eggs, which were 80—90 um in diameter. Larvae hatched with statocysts and a distinct opercu- lum but without eyes. A pigmented mantle organ is evident from Figure 32 in Ostergaard (1950). No other observations of the embryology or larval type have been published for the entire superfamily. Here I describe the embryology and larval development of Tylodina fungina as a step towards documenting development in this phylogenetically important group. MATERIALS AND METHODS 12 adult Ty/odina fungina were collected by dredging in the Perlas Isands (8°35.9'N, 78°1.0’W and 8°16.0N, 79°1.3W) during February and April 2007. The snails were brought to the surface attached to the host sponge Page 52 the Veliger, Vols 50Noml Suberea azteca (Gomez and Bakus, 1992). Tylodina fungina is usually reported associated with Ap/ysina fistularis, which appears superficially similar to S. azteca. Identification of S. azteca was verified from a preparation of skeletal material and comparison with the original species description. The snails and some host sponge were kept in running seawater at ambient temperature (22—26°C). The sponge survived for 2 weeks under these conditions, but the snails survived for up to 6 weeks. Portions of egg ribbon were scraped from the surface of the containers and collected from the sponge skeletons and maintained in fingerbowls in the laboratory at 21—23°C. The water was changed daily and larvae were collected immediately upon hatching. After hatching the larvae were transferred to finger bowls with | um filtered water. The water was changed every 2—3 days and larvae were fed [sochrysis galbana. The hydrophobic larvae were kept from getting stuck in the surface tension of the water by the addition of a few flakes of cetyl alcohol. Only uncleaved eggs and naturally hatched larvae were measured. RESULTS In the laboratory, the adult Ty/odina fungina remained closely associated with the live sponge and were frequently observed feeding on it (Figure 1A). One of the sponges had been completely consumed by the snails and all that remained was the spongin skeleton. This skeleton was covered with egg ribbons (Fig- ure 1B), giving the appearance of badly damaged sponge tissue when, actually, no sponge tissue remained on the skeleton. The three other sponges that remained largely intact showed eroded areas which each housed a snail (Figure 1A). Egg ribbons were not evident on these sponges, which suggests that egg production commences after depletion of the food supply. After the sponges died the snails deposited egg ribbons on the containers 1n which they were housed (Figure 1C). The bright yellow egg ribbons were attached flat against the substrate and were arranged in an irregular spiral when laid on a smooth surface (Figure IC). Those that were attached to the sponge skeleton were irregularly twined around the skeleton and incorporat- ed portions of the skeletal fibers (Figure 2A). The 80.5 um (n = 29, s. d. = 1.4 um) eggs were yellowish cream-colored and were each contained within a 125.1 um capsule (n = 19; s.d. 3.5 um). These capsules are embedded in rows within the gel of the egg ribbon. Between the rows of egg capsules there were bright yellow streaks of extracapsular material (Figure 2 B— F). These streaks were inconsistent in width and were absent from some portions of the egg ribbon, but when present there tended to be 2 rows of eggs between each streak (Figure 2). At high magnification the streaks Table 1 Developmental time table of Tylodina fungina at 2\— 24°C. Time Stage 24 hrs Blastula 48 hrs Gastrula 4 days “Trochophore”’ 6 days Shell begins to develop 7 days PMO develops 13-15 days Hatching could be seen to consist of numerous tiny droplets (Figure 2F), which remained in the gel after hatching. Several egg ribbons were collected prior to first cleavage and were observed until hatching. A develop- mental schedule is given in Table 1. The two polar bodies remain associated with the eggs at least until gastrulation (Figure 3D). The first two cleavages appear to be equal and synchronous and there is no polar lobe (Figure 3B, C). By the beginning of the third cleavage division, one of the 4 cells is already slightly ahead of the others. Later, cleavage becomes more asynchronous and eventually forms a compact, animal- vegetally flattened blastula (Figure 3D). The gastrula is horseshoe shaped and appears to have been formed at least partially by invagination (Figure 3E). A trocho- phore-like stage with a distinct raised ring of cilia around the anterior end (Figure 3F) follows gastrula- tion. The pre-hatching veliger shows a distinct foot with an operculum (Figure 3G) and pair of statocysts and a large, pigmented mantle organ (PMO) on the right side (Figure 3G, H). The PMO appears black with epi-illumination and is dark red under transmitted light. At hatching the larvae have a round, transparent shell 123.1 um (n = 58 from 3 ribbons; s. d. = 6.1) in length with a single slightly left-handed whorl. On living larvae the shell appears smooth, but slight granular sculpture is evident on dead shells. The velum is un-pigmented and consists of two small, equal, semicircular lobes (Figure 3H). The operculum is present and the foot is simple. After 3 weeks the larvae had grown to 161.8 um (n = 7; s. d. = 8.3) but still had not developed eyes or tentacles and showed no signs of competency to settle. The larvae survived for at most 4 weeks in culture. Despite repeated attempts to culture them, it was not clear why they failed to thrive. DISCUSSION As previously noted by Robertson (1985), develop- mental features have the potential to contribute useful data to understand high-level gastropod relationships. The main drawback to using developmental features is R. Collin, 2007 Page 53 Figure 1. Adult 7. fungina with host sponge and egg masses. A. Two adult T. fungina on their host sponge. The smaller individual (arrow) is sheltered in a depression in the ectosome of the sponge. Scale = 4 cm; B. Skeleton of the host sponge covered with egg masses of 7. fungina. Scale = 2 cm; C. Egg mass of 7. fungina deposited on a plastic mesh. This mass is smaller and more tightly coiled than most of the masses deposited on flat surfaces. Scale that few data are available for many interesting groups. Tylodinoids are a prime example of a phylogenetically important group where little is known. However, some comparisons of the egg masses can be made with previously published observations. The egg masses of T. fungina as described here seem generally similar to those of the Australian congener, T. corticalis, with a flat ribbon attached in a coil to the substrate. Egg masses of both species are yellow, but 1 cm. contain cream-colored eggs (Thompson, 1970), sug- gesting that T. corticalis, like T. fungina, deposits extracapsular material in the egg ribbons. The differ- ence in egg size between the 80 um eggs of T. fungina and the 98 um eggs of 7. corticalis further bolsters their status as distinct species. The lack of information on Umbraculum species makes it difficult to determine how consistent the egg masses are throughout the family. The large number of eggs per capsule in Umbraculum the Veliger, Vol, s03Nomi Figure 2. Egg ribbons of T. fungina. A. Egg ribbon attached to the sponge skeleton; Scale = 1 cm; B. Egg ribbon that was laid on a smooth surface, the lines of extra-capsular material are clearly visible in color but difficult to see in black and white. Scale = | cm; C. The multiple layers of eggs and the uneven distribution of the extracapsular material are can be seen. Scale = 5 mm; D. and E. Closer views showing the arrangement of eggs in two rows between each string of opaque extracapsular material. Scale = 1.5 mm and 500 um respectively; F. Detailed view of egg capsules embedded in the gel and the droplets of extracapsular material. These droplets remain in the gel after hatching. Scale = 150 um. sinicum (Thompson, 1970) does show that there are some differences. Unlike the Tylodinoids, there is considerable pub- lished information on the development of the other notaspidean superfamily, the Pleurobranchoidea (re- viewed in Gibson, 2003). Gibson (2003) described the typical features of notaspidean development on the basis of her detailed observations of the development of Pleurobranchaea maculata and a review of the litera- ture. These new observations of Ty/lodina development suggest that tylodinid development may differ signifi- cantly from pleurobranchid development. Unlike pleurobranchids, tylodinids have a larval operculum and extracapsular material (Table 2). Unfortunately, the larvae in this study did not survive long enough to ermine if the larval mantle overgrows the larval shell unusual characteristic of pleurobranchids). It is unlikely, however, that this would happen as adult Tylodinids, unlike pleurobranchids, have a fairly large external shell that is not covered by the mantle. It may be that the mantle overgrowth of the larval shell is what prevents pleurobranchid larvae from being hydropho- bic, like other opisthobranch veligers. The most unusual characteristic of the Ty/lodina fungina egg masses was the presence of extracapsular material. Similar material in opisthobranch egg ribbons is usually referred to as “yolk” in the literature, although there is usually little evidence beyond a similar color that suggests this material is indeed yolk. “Yolk bodies” embedded in the egg ribbon jelly outside the egg capsules are well known for tropical chromo- dorids and sacoglossans (Boucher, 1983). Boucher (1983) described three kinds of extra-capsular material. Chromodorids have yolk that is present as either cap- R. Collin, 2007 Page 55 Figure 3. Developmental stages of T: fungina. A. Uncleaved eggs individually enclosed in transparent capsules, which have been removed from the gelatinous mass. B. 2-cell stage, the polar bodies are not visible. C. 4-cell stage. D. The flattened blastula with the polar bodies marking the animal pole. E. Horse-shoe shaped gastrula F. Late ““trochophore’’stage with the beginnings of the shell showing on the bottom right G. Encapsulated veliger, 2 days prior to hatching. The red PMO 1s visible on the animal’s right and has not yet developed the black pigmentation. The velum, foot, and operculum are all well developed. H. 2-week old veliger larva. Scale = 100 um. like “yolk bodies” associated with individual capsules or discrete “‘yolk”’ masses distributed through the egg mass. Elysia species have strings of “yolk” running through the egg masses (Boucher, 1983). The overall morphology of Elysia egg masses is strikingly similar to those described here for 7. fungina (P. Krug, pers. com). It has yet to be determined if the material included in the T. fungina egg masses is yolk, but it seems unlikely. The material is a different color (bright yellow) from the eggs (cream) and remains in the gel of the egg mass after hatching. The presence of this “yolk” in several other species with planktotrophic development, where the larvae are not retained near the egg mass after hatching (Boucher, 1983) suggests that this material might not have a nutritive function. There iS some circumstantial evidence that the function in Tylodina might be defensive. Becerro et al. (2003) showed that egg masses and extracts of egg masses from Tylodina perversa deter feeding by damselfish with the same efficiency as the chemically defended adult snails and Ebel et al. (1999) showed that defensive chemicals are sequestered in the egg masses of the same species. Detailed examination of this material is necessary before their function can be determined. Acknowledgments. I thank the captain and crew of the R/V Urraca without whom I would not have collected the samples, A. Baeza for photographing the adult animals, M. C. Diaz and R. Thacker for identifying the sponge, and the Autoridad Maritima de Panama for issuing necessary permits to the Smithsonian Tropical Research Institute. Table 2 Comparisons of Tylodinid and Pleurobranchid development. Character Tylodinids Pleurobranchids Egg masses Flat ribbons Strings Extra-capsular material Present Absent Extra-embryonic, intra-capsular yolk Absent Present sometimes Type | larval shell Present Present Larval Shell Hydrophobic Not hydrophobic in Pleurobranchaea Larval shell growth Operculum Present PMO Present Larval eyes Absent at hatching No observations of mantle-over growth * maculata ¢ Over-grown by mantle Absent 8 Present Absent at hatching * Glenys Gibson, pers. com. 2007 * More data is necessary to verify this observation. * Reported as absent in the group by Gibson (2003), but curiously Ostergaard (1950) reported opercula on 2 species of Pleurobranchids. Opercula were not present in other published studies of development in this group. Page 56 LITERATURE CITED BECERRO, M. A., X. TURON, M. J. URIZ & J. TEMPLADO. 2003. Can a sponge feeder be a herbivore? Ty/odina perversa (Gastropoda) feeding on Aplysina aerophoba (Demospongiae). Biological Journal of the Linnean Society 78:429-438. BOUCHER, L. M. 1983. Extra-capsular yolk bodies in the egg masses of some tropical opisthobranchia. Journal of Molluscan Studies 49:232—241. EBEL, R., A. MARIN & P. PROKSCH. 1999. Organ-specific distribution of dietary alkaloids in the marine opistho- branch Tylodina perversa. Biochemical Systematics and Ecology 27:769-777. Gasp, W. M. 1865. Description of new species of marine shells from the coast of California. Proceedings of the California Academy of Science 3:182—190. GIBSON, G. D. 2003. Larval development and metamorphosis in Pleurobranchaea maculata, with a review of develop- ment in the Notaspidea (Opisthobranchia). Biological Bulletin 205:121—132. The Veliger, Vol. 50, No. 1 OSTERGAARD, J. M. Spawning and development of some Hawaiian marine gastropods. Pacific Science, 4:75—115. ROBERTSON, R. 1985. Four characters and the higher category systematics of gastropods. American Malacological Bul- letin, Special Edition 1:1—22. THOMPSON, T. E. 1970. Eastern Australian Pleurobrancho- morpha (Gastopoda: Opisthobranchia). Journal of Zool- ogy, London 160:173-198. WAGELE, H. & R. WILLAN. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the Linnean Society 130:83-181. WILLAN, R. C. 1987. Phylogenetic systematics of Notas- pidea (Opisthobranchia) with reappraisal of families and genera. American Malacological Bulletin 5: 215-241. WILLAN, R. C. 1998. Order Notaspidea. Pp. 977-980 in P. L. Beesley, G. J. B. Ross & A. Wells (eds.), Mollusca: The Southern Synthesis. Fauna of Australia. Vol. 5. CSIRO Publishing: Melbourne. Part A xvi 563pp. The Veliger 50(1):57 (March 11, 2008) THE VELIGER © CMS, Inc., 2007 BOOK REVIEW The Recent Molluscan Fauna of fle Clipperton (Tropical Eastern Pacific) by KIRSTIE L. KAISER. Festivus 39, Supplement. 162 pp. The atoll of Clipperton is not only the most isolated island in the tropical eastern Pacific, but its marine fauna is a unique biogeographic mixture of species, with approximately equal parts arriving from the Indo- West Pacific and Panamic Provinces. Kirstie Kaiser’s new volume on the molluscs of Clipperton is a substantial and welcome addition to the study of this little-known fauna. Some 285 molluscan species, including planktonic species and two terrestrial gastro- pods, are now known from the atoll, a huge increase from the 92 species recorded previously. The volume contains illustrations, details of collecting localities, a list of recorded species and their geographic distribu- tion, and an account of rejected records. A striking feature of this fauna is the large number of minute species, many of which probably have brief pelagic larval stages or none at all. Unfortunately, most of these taxa are identified only to genus or even family level, so that their phylogenetic and biogeographic affinities remain unknown. Some major tropical groups are entirely absent in Clipperton’s fauna. For example, there are no patello- gastropod limpets, turban snails (Turbininae), strom- bids, olives (Olividae), siphonariid (or pulmonate) limpets, cockles (Cardiidae), venerids, and _tellinids. Although many of these groups comprise sand-dwellers, and sand is rare at Clipperton, some sand-dwelling gastropods do occur on the island, including a few cones, terebrids, moon snails (naticids), and a nassariid. These faunal peculiarities raise interesting questions that for the most part are neither posed nor discussed in this volume. Taxonomic remarks are given for some species, including members of the bivalve genus Chama, but for most species such annotations are wanting. It would have been interesting, for example, to know why Kaiser recognizes two species of the coral-associated muricid genus Quoyula (or, perhaps more properly, Galeropsis) instead of the single species that most recent authors recognize. The Panamic muricid genus Plicopurpura is represented at Clipperton by the form identified by Kaiser as P. pansa (Gould, 1853). The name P. pansa refers to the broad-apertured morph of a_ highly variable species that is usually known by its senior synonym, P. columellaris (Lamarck, 1816). The name and the illustration imply that only the broad- apertured form of this species is present at Clipperton, a situation parallel to that of the sister species P. patula (Linnaeus, 1758) in the West Indies; and that the narrow-apertured, thick-shelled morph with strong outer-lip teeth, to which the name P. columellaris was initially applied, would appear to be absent on Clipperton. Is this so? An important question about island faunas is whether the species found there maintain viable populations. A virtue of Kaiser’s compilation is that dates of collection are given for each species, together with whether specimens were taken dead or alive. Kaiser implies that the two species of the lucinid bivalve genus Codakia at Clipperton may no longer be living in the lagoon, as they did when the lagoon was open to the ocean. Many other species are also recorded only as empty shells, implying that many taxa recorded from the atoll are only occasionally present there. Although it is obvious that a great deal of basic taxonomic work remains to be done, Kaiser’s volume offers a solid foundation on which further studies can be built. Questions about island diversity, interactions between species originating in different biogeographic settings, sustainability of island populations, and niche shifts made possible by the absence of major predators and competitors cannot be answered without descrip- tive faunal accounts of the kind Kaiser has brought together for Clipperton. Geerat J. Vermeij i scanuigtt ul is te 4 Instructions to Authors The Veliger publishes original papers on any aspect of malacology. All authors bear full responsibility for the accuracy and originality of their papers. Presentation Papers should include an Abstract (approximately 5% of the length of the manuscript), Introduction, Materials and Methods, Results, and Discussion. Short Notes should include a one- sentence Abstract. In taxonomic papers, all names of taxa must be accompanied by author and date of publication, and by a full citation in the bibliography. In papers on other subjects and in the non-taxonomic portions of taxonomic papers, author and date of names need not be accompanied by a full citation. All genus and species names should be in italics. All references to new molecular sequences must be linked to GenBank. Literature Cited Each citation should consist of a complete list of authors, with initials, date of publication, full title of article of chapter, names of all editors of books, publisher and place of publication for all books, journal titles spelled out in full, volume number (and, where appropriate, issue number), and inclusive pagination. Authors must be consistent about punctuation. Every reference in the text must be accompanied by a citation in the bibliography, and all entries in the bibliography must be mentioned in the body of the paper. Authors should cross-check the bibliography with the text. Submitting manuscripts All manuscripts should be submitted as Word files, by e-mail or on disc, double spaced, plus one optional double-spaced paper copy. For convenience of review, figures are best submitted initially in a form and size suitable for electronic mail. Figures for publication may be submitted on disc. Halftones should be at least 300 ppi; graphics in TIFF or EPS format. Send manuscripts, proofs, books for review, and correspondence on editorial matters to: Geerat J. Vermeij Editor, The Veliger Department of Geology University of California at Davis One Shields Avenue Davis, CA 95616 veliger@geology.ucdavis.edu T 530.752.2234 F 530.752.0951 In the cover letter, authors should briefly state the point of the paper, and provide full and electronic addresses of at least three reviewers who have not previously seen the manuscript. If authors feel strongly that certain reviewers would be inappropriate, they should indicate reasons for their views. CONTENTS — Continued BOOK REVIEW The Recent Molluscan Fauna of Ile Clipperton (Tropical Eastern Pacific). Kirstie L. KAISER GEERAT J, VERMEIL. 2222 sa cetes 32 oe oe eae a eens ei Rae tia tote DY, SMITHSONIAN eee LIBRARIES IAEA 3 9088 01431 8661 (HE me LICER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler (1901—2000), Founding Editor Volume 50 L (a UX AOLL— ISSN 0042-3211 June 20, 2008 Number 2 CONTENTS Synecology of a Springsnail (Caenogastropoda: Hydrobiidae) Assemblage in a Western U.S. Thermal Spring Province 1D) OAT NAR SAID An cae ye pesrcieetels seeeu select nee) legion eee pcan atinnsee WN ah aren tart tty tia a uel 59 Two New Species of the Genus Cerithiopsis Forbes & Hanley, 1850 (Gastropoda: Cerithiopsidae) from Brazil RaQueEL MEDEIROS ANDRADE FIGUEIRA AND ALEXANDRE Dias PIMENTA.............45 Yo A Record of the Invasive Slug Veronicella cubensis (Pfeiffer, 1840) in California ReWwVic DONNELIT, A. EIANSEN, ID: PAINE ANDi M: J. GORMALLY ..........2.-¢..--% 81 Developmental Mode in Opisthobranch Molluscs from the Tropical Eastern Pacific Ocean JEEEREY@ bl Re GODDARD AND ATACIATIERMOSILEO!: «244s sia eae seine meres ceo 83 Three New Buccinid Species (Gastropoda: Neogastropoda) from Chilean Deep-Water, Including One from a Methane Seep INOENGERAUSSENPANIDEAVIERE SETI ANIES S04 yanie i) secu eau allie acy gelee Cyn coe ea lem seu i Redescription of the Deep-sea Wood Borer Neoxylophaga teramachii Vaki & Habe, 1950 and its Assignment to the Genus Xyloredo (Bivalvia: Myoida: Pholadoidea) with Comments on Fossil Pholadoidae TOAUINIA JELACA VANS) IOIMCONSIIKAG GS ucaieis pode ob coco moooben boule ude Go bU cede con Our 107 A Note on Strombus coronatus Defrance, 1827 and Strombus coronatus Réding, 1798 (Mollusca: Gastropoda) Marutas HaRZHAUSER AND Gis C. KRONENBERG 1.0... ccc ee eee eee 120 CONTENTS — Continued The Veliger (ISSN 0042-3211) is published quarterly by the California Malacozoological So- ciety, 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 His- tory, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. “OTHE WELIGER >: Scope of the Journal | The Veliger is an international, peer-réviewed scientific quarterly published by the California Malaco- zoological Society, a non-profit educational organization. The Veliger is open to original papers pertain- ing to any problem connected with mollusks. Manuscripts are considered on the understanding that their contents have not appeared, or will not appear, elsewhere in substantially the same or abbreviated form. Holotypes of new species must be deposited in a recognized public museum, with catalogue num- bers provided. Even for non-taxonomic papers, placement of voucher specimens in a museum is strongly encouraged and may be required. Editor-in-Chief Geerat J. Vermeij, Department of Geology, University of California at Davis, One Shields Avenue, Davis, CA 95616 e-mail: veliger@geology.ucdavis.edu Managing Editor Edith Vermeij Board of Directors Terrence M. Gosliner, California Academy of Sciences, San Francisco (President) Hans Bertsch, Tijuana and Imperial Beach Henry W. Chaney, Santa Barbara Museum of Natural History Matthew J. James, Sonoma State University Rebecca F. Johnson, California Academy of Sciences, San Francisco Michael G. Kellogg, City and County of San Francisco Christopher L. Kitting, California State University, Hayward David R. Lindberg, University of California, Berkeley Peter Roopharine, California Academy of Sciences Barry Roth, San Francisco Angel Valdés, Natural History Museum of Los Angeles County Geerat J. Vermeij, University of California, Davis Membership and Subscription Affiliate membership in the California Malacozoological Society is open to persons (not institutions) interested in any aspect of malacology. New members join the society by subscribing to The Véeliger. Rates for Volume 50 are US $65.00 for affiliate members in North America (USA, Canada, and Mexico) and US $120.00 for libraries and other institutions. Rates to members outside of North America are US $75.00 and US $130.00 for libraries and other institutions. All rates include postage, by air to addresses outside of North America. Memberships and subscriptions are by Volume only. Payment should be made in advance, in US Dollars, using checks drawn from US banks or by international postal order. No credit cards are ac- cepted. Payment should be made to The Veliger or “CMS, Inc.” and not the Santa Barbara Museum of Natural History. Single copies of an issue are US $30.00, postage included. A limited number of back issues are available. Send all business correspondence, including subscription orders, membership applications, payments, and changes of address, to: The Veliger, Dr. Henry Chaney, Secretary, Santa Barbara Museum of Natural His- tory, 2559 Puesta del Sol Road, Santa Barbara, CA 93105, USA. Send manuscripts, proofs, books for review, and correspondence regarding editorial matters to: Geerat Vermeij, Department of Geology, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). (JL 072008) ~ LIBRARIES THE VELIGER z The Veliger 50(2):59-71 (June 20, 2008) © CMS, Inc., 2007 Synecology of a Springsnail (Caenogastropoda: Hydrobiidae) Assemblage in a Western U.S. Thermal Spring Province DONALD W. SADA Division of Hydrologic Sciences, Desert Research Institute, 2215 Raggio Parkway, Reno, NV 89512, USA (e-mail: Don.Sada@dri.edu), fax: 775/673-7363 Abstract. Springsnails are numerically dominant members of benthic communities in many springs of western North America and Australia. Several studies have shown the influence of water chemistry on their abundance and distribution within springs, but little information exists regarding their use of physical aspects of spring environments. Habitat preferences, niche breadth and overlap, and environmental factors influencing the structure of an assemblage of native springsnails (Pyrgulopsis avernalis, P. carinifera, and Tryonia clathrata) and the non- native red-rimmed melania (Melanoides turberculata) gastropod are described in a southern Nevada, USA, thermal spring province. Water temperature, current velocity, and substrate type were the most important physical factors structuring the assemblage. Springbrook wetted width, presence of armored and incised banks, and location of sample sites across the wetted width were also statistically significant, but less important factors. Each species occupied a wide diversity of habitats, but each species also exhibited habitat preferences for a range of depths, velocities, temperatures, or substrates. Niche overlap varied among species and habitats were partitioned among species for a minimum of two environmental resources. Competitive interactions appeared to minimally influence the structure and distributions of species belonging to this assemblage. Findings suggest that springsnails are restricted to portions of a spring that provide suitable physicochemical conditions, and that each springsnail taxon may exhibit specific habitat requirements. Springsnail extinctions and declines in abundance in western North America and Australia can be attributed to human activities altering the discharge and water depth, substrate composition, current velocity, and water temperature of springs. Novel approaches are required to alter human uses and facilitate restoration, and protect the integrity of these unique arid land aquatic systems. Key Words: Spring ecology, Hydrobiidae, springsnail ecology, arid land wetlands. INTRODUCTION The Hydrobiidae is a worldwide family of primarily freshwater gill-breathing gastropods. Recent taxonom- ic studies have found an amazing diversity of hydro- biids (commonly referred to as springsnails) in isolated, arid land springs of North America and Australia. More than 120 species in seven genera are known from more than 1000 springs in the western U.S., and 35 species in nine genera from Australia (Ponder et al., 1989; Hershler, 1994, 1998, 2001). Springsnails are restricted to persistent aquatic habitats that are minimally affected by drought (Taylor, 1985) and most species occupy few localized habitats within a limited studies showing the influence of temperature on Pyrgulopsis bruneauensis demography and feeding in springs along the Snake River of southern Idaho (Mladenka and Minshall, 2001), and relationships between CO, concentrations and P. montezumensis abundance in northern Arizona (O’Brien and Blinn, 1999). Richards et al. (2001) examined spatial relation- ships in an assemblage of three snails including Taylorconcha serpenticola and Ponder et al. (1989) examined the influence of several environmental factors on activity and survivorship of several species in mound springs of Australia. These species responded to desiccation, salinity, deoxygenated water, water temperature, and submersion but the authors were geographic area (Hershler, 1998). They are usually the most abundant benthic macroinvertebrate in springs where they occur, and springs occupied by more than one springsnail species are uncommon. In spite of their taxonomic diversity and _ their abundance in springs, few ecological studies have been conducted. Information is limited to demographic unable to quantify relationships between habitat zones and springsnail abundance. Qualitative observations during taxonomic and biogeographic surveys suggested that abundance within a spring is influenced by water depth, current velocity, substrate composition, and aquatic vegetation (e.g., Hershler and Sada, 1987; Hershler, 1998). These observations indicate that each Page 60 The Veliger) Vol) 50> New species occupies a unique habitat within a spring, but they have not been supported by quantitative studies examining relationships between spring environment, microhabitat use, and assemblage structure. Ecological information is needed to provide insight into isolating mechanisms that have facilitated development of this diverse fauna in small, isolated habitats. This informa- tion is also important in determining how springsnails respond to human activities that alter springs. These small wetlands support much of the aquatic life in arid lands and most springs have been degraded by livestock, diversion, and introduction of non-native species (Sada et al., 1992; Shepherd, 1993; Hubbs, 1995; Myers and Resh, 1999; Sada and Vinyard, 2002; Sada et al., 2005). These activities have justified listing several springsnail species as endangered in the western U.S., and Hershler (1998) and Sada (field notes) recorded three extinctions and extirpation of 13 populations in the past decade. In this study, an assemblage of four species (three springsnails and one non-native mollusk) was exam- ined in a thermal spring province to: 1—determine physical factors (in addition to temperature) affecting assemblage structure, 2—quantify microhabitat use, and 3—determine if habitat use differs among species. The spring province includes approximately 30 springs that discharge a total of 1.3 m*/sec. Discharge and temperature at individual springs ranges from 10— 200 I/min, and 24.5—31.8°C, respectively. This assem- blage included Pyrgulopsis avernalis and P. carinifea that are endemic to this province, and Tryonia clathrata that is endemic to thermal springs along the pluvial White River system in eastern Nevada (Hersh- ler, 1994, 2001). All of these species are small. The shell height of P. avernalis ranges from 2.4 mm to 4.3 mm, for P. carinifera from 3.8 mm to 5.0 mm, and for 7. clathrata from 2.9mm to 7.0 mm. The red-rimmed melania (Melanoides tuberculata, family Thiaridae), which is native to Asia (Burch and Tottenham, 1980), also inhabits these springs. Shell height of adult melania is from 1 cm to 3 cm. Quantitative studies have not assessed the influence of this species on springsnails. Hershler and Sada (1987) noted that springsnail abundance may be decreased in its pres- ence, and Pointier et al. (1993) and De Marco (1999) found that it detrimentally affected native gastropod abundance. This work was one component of studies examining the benthic macroinvertebrate communities in this spring province, which will be the subject of a subsequent article. SITE DESCRIPTION Studies were conducted in a spring province (collec- tively referred to as ‘Warm Springs’) that forms the Muddy River in Clark County, Nevada (Figure 1). The Muddy River flows approximately 35 km into the Colorado River (which is now within the Overton Arm of Lake Mead). The springs creating the Muddy River are located at approximately 500m elevation and scattered over approximately 2,000 hectares. Water flows through approximately 4 km of springbrook before forming the Muddy River. Water temperature at spring sources is approximately 32°C, and combined discharge from the province is a relatively constant 1.5 m*/sec (Eakin, 1964). Discharge from individual springs ranges from approximately 0.0028 to 0.17 m?/ sec and spring brooks are bordered by ash (Fraxinus velutina), mesquite (Prosopis sp.), non-native salt cedar (Tamarisk sp.), and fan palm (Washingtonia filifera). These woody species are interspersed with grasses (mostly Distichlis spicata) and perennial herbs. Spring- brooks support diverse aquatic habitats from low gradient brooks that meander over fine substrates to higher gradient brooks where swift water flows over gravel and cobble substrates. General characteristics of spring brooks that were sampled during this study are summarized in Tables | and 2. In addition to mollusks, these springs support a number of rare fishes and other aquatic macroinvertebrates, many of which are endem- ic to Warm Springs (U.S. Fish and Wildlife Service, 1996, Hershler, 1994, 2001; Schmude, 1999; Polhemus and Polhemus, 2002; Sada and Vinyard, 2002). Native Americans had settlements near Warm Springs, and in the early 19th Century it was settled by white men. Its springs have been altered for recreation and diversion, channelization, and siltation from agriculture, and non-native fishes and aquatic invertebrates have been introduced (Scoppettone, 1993). These alterations reduced native fish abundance and resulted in listing the Moapa dace (Moapa coriacea) as endangered by the U.S. Fish and Wildlife Service (U.S. Fish and Wildlife Service, 1996). Since the early 1980s, Moapa dace recovery programs have restored six springs and approximately 600 m of spring brook (approximately 15 percent) to natural condition. METHODS Data Collection Aquatic habitat parameters (Tables 1 and 2) and mollusks were sampled in 84, 10 cm X 12 cm quadrats at 21 stations located at predetermined distances from five spring sources (two with long and three with short spring brooks) (Table 3) during the autumn of 1996. Sample stations included the diversity of habitats in first-order spring brooks, and were placed near springs sources and along the downstream continuum to the confluence of the nearest tributary spring brook. The distance of stations from spring sources varied because D. W. Sada, 2007 Page 61 Baldwin Springs Apcar Springs Ov Pedersen Springs oo Plummer Springs * Sample point o& Spring Figure 1. Map showing the approximate location of sample sites (*) and major springs in the ‘Warm Springs’ province that combine to form the Muddy River, Clark County, Nevada. some reaches were not easily accessed, some spring brooks were less than 50 m long, and longer intervals occurred in large springs with long spring brooks. Each station consisted of four transects spanning the wetted Table 1 Median and range of aquatic habitat parameters measured (units shown in parentheses) within quadrats and at 21 stations where mollusks were sampled in Warm Springs, upper Muddy River. All variables used in CCA to examine relationship between environmental factors and mollusk assemblage structure. * = parameters measured in quadrats, ** = parameters measured along transects, all others measured at stations. Element Median Range Water Depth (cm)* 16 2-70 Water Velocity (cm/sec)* 20 0-109 Springbrook Width (cm)** 180 40-530 Water Temperature (°C) 31.3 24.5-31.8 Dissolved Oxygen (mg/l) Dull 4.15.7 Electrical Conductance 1100 1050-1100 (umhos/cm) pH 7.6 7.4-7.6 Riparian Cover (percent)** 90 2-100 width (spaced 1 m apart and oriented perpendicular to the thalweg) where mollusk samples and aquatic habitat measurements were collected from two mid- channel and two springbrook margin quadrats. Mid- channel quadrats were placed along first and third Table 2 Proportion of quadrats and spring brook banks where substrate and channel features, respectively, occurred during mollusk sampling in Warm Springs, upper Muddy River, Clark County, Nevada. n = 84 for Substrate Features (all measured in quadrats) and n = 168 for Channel Features (all measured where transects intersected banks). All variables used in CCA. Substrate Channel Feature Proportion Feature Proportion Fines 0.28 Stable Channel 0.87 Sand 0.18 Incised Banks OME Gravel 0.44 Bank Overhang 0.57 Cobble 0.11 Bank Perennial 0.42 Vegetation CPOM 0.16 Grassy Banks 0.14 Palm Roots 0.43 Armored Banks 0.14 Page 62 Table 3 Spring brook names and the approximate distance of sample stations from each spring source. Springbrook Name Locations (m) Pedersen Spring 10, 25, 60, 120, 200, 500 Muddy Spring 10, 25, 60, 120, 180, 280 Plummer Spring 10, 25, 60, 120 Apcar Spring 25, 60 South Fork Muddy River Spring — 25, 60, 120 transects, and right and left bank quadrats on transects two and four, respectively (bank quadrats scored 1 and mid-channel scored 0 for canonical correspondence analysis). Channel features (Table 2) and springbrook width were recorded across each transect and bank features were recorded where transects intersected the banks. Water depth and mean water column velocity (measurement taken at 60 percent water depth) were measured at the center of each quadrat, and occurrence of substrate types, filamentous green algae, palm roots, and coarse particulate organic matter (CPOM) were scored as present (1) or absent (0) from a quadrat. Electrical conductance (EC), dissolved oxygen concen- tration, temperature, and pH were measured at each station (using Model 33 [EC and temperature] and Model 57 YSI [dissolved oxygen] meters, and an Oakton pHTestr 2 handheld meter). A Marsh-McBir- ney Model 2000 current meter was used to measure current velocity. Although this velocity measurement may weakly quantify benthic microhabitats, correlation between these velocities suggests that conditions at mean water column are indicative of velocities over substrate. pH and dissolved oxygen meters were calibrated daily and other meters calibrated according to manufacturer specifications. Mollusks were collected by roiling substrate within the quadrat for 10 sec to flush material downstream into a 250 micron mesh net that was held in a vertical frame and secured to the quadrat. Samples were preserved in 90 percent ethyl alcohol and returned to the laboratory for identifica- tion, and enumeration. Identification was made using descriptions in Hershler (1994, 2001). Samples are archived at the Desert Research Institute, Aquatic Ecology Laboratory, Reno, Nevada. Data Analysis Environment-Assemblage Relationships: Relationships between aquatic and channel environments and assem- blage structure were examined with canonical corre- spondence analysis (CCA) using Canoco 4.0 for Windows. CCA axis scores were standardized using methods of Hill (1979), scaled to optimize the representation of species, and Monte Carlo simulation The Veliges-Volks0s Now (1000 iterations) tested the hypothesis that there was no relationship between species and environment matrices. A total of 22 measured and categorical habitat features were included in the analysis (EC and pH did not differ among stations and were not used in the CCA). The CCA is a multivariate direct gradient analysis that analyzes unimodal data to assess species distribution along environmental gradients. It performs multiple linear least-squares regressions with the environment and species abundance as independent and dependent variables, respectively (ter Braak and Prentice, 1988; Jongman et al., 1987; Palmer, 1993). Microhabitat Use: Habitat preference was calculated for water depth, temperature, and velocity, and the presence of substrate types. With the exception of water depth, CCA showed these variables were most important to structuring the molluscan assemblage (Figure 2, Ta- ble 4). Preferences for water depth were calculated because preliminary analysis indicated these species occupied a diversity of available depths. Preference was calculated using the formula of Jacobs (1974): D = r — p/ r +p — 2rp; where p is the proportion of the resource available in the habitat and r is the proportion of the resource utilized by the species. Resource use was categorized as moderate preference (between 0.25 and 0.5) or strong preference (>0.5), or strongly (<—0.5) or weakly avoided (between —0.25 and —0.5). Neither preference nor avoidance were indicated by values < 0.24 and > —0.24. Niche breadth was calculated using the equation B = 1/9’ P;;*, where Pj; is the proportion of the resource in each category (Levins, 1968), and niche overlap was calculated using the same variables follow- ing the equation from Schoener (1970) with revisions by Litton et al. (1981) so that S = 100(1—1/2 )|Pxi — Pyil) where P,; and Py; are the proportion of resource use in each category for the two species being compared. Niche breadth values may range from 1 — > 14. Low values are indicative of narrow, limited habitat use and high values indicate wider use of available habitats. Niche overlap values range from 0 to 1 with substantial overlap being indicated by values > 0.5 and differences in use indicated by values < 0.5. Habitat availability was calculated using records from all quadrats (n = 84) and habitat use, and niche breadth and overlap calculations were made by weighting resource utilization in accordance with each species’ abundance. Preference, niche breadth, and niche overlap calculations must be interpreted with caution. They are indices that provide insight into relationships in habitat use among species but they are not quantitative descriptions of inter-specific interactions, which can only be determined through experimental manipulation. Similarities among results of these analyses and CCA may guide experimental studies by indicating the relative influence of individual environmental variables on the distribution of species. D. W. Sada, 2007 +2.0 Axis 2 -2.0 Page 63 MT i +2.0 +2.0 Axis 1 Figure 2. Canonical correspondence analysis biplot showing the relative influence of significant instream and channel environmental parameters on structure of the mollusk assemblage at Warm Springs. The relative influence of each environmental variable is shown by vector length with longer vectors indicating variables with greater importance. PA = P. avernalis, PC = P. carinifera, TC = T. clathrata, MT = M. tuberculata, WV = mean water column velocity, TEMP = water temperature, GR = gravel, FINES = fines, IB = presence of incised banks, POS = quadrat position, CPOM = coarse particulate organic matter, WW = wetted (spring brook) width, and BA = presence of armored bank. RESULTS Environment-Assemblage Relationships: A total of 1282 P. carinifera, 704 P. avernalis, 750 T. clathrata, and 283 M. tuberculata were collected and tallied. Snail abundance in quadrats ranged from 0 to 346 (equiv- alent to approximately 29,000/m?), and from 0 to 148 for P. carinifera, 0 to 145 for P. avernalis, 0 to 96 for T. clathrata, and 0 to 34 for M. tuberculata. Initial CCA revealed nine significant (P < 0.05) physicochemical factors were most important to structuring the assemblage (Table 4). Water temperature, mean water column velocity, the presence of some substrates (fines, gravel, CPOM), aspects of channel morphology (incised or armored banks, spring brook width), and quadrat location (bank vs. mid-channel) were statisti- cally significant variables. Water depth, the presence of roots, algae, other submerged vegetation, sand and cobble substrates, dissolved oxygen concentration, distance from spring source, and bank angle, stability and grassy vegetation had comparatively little influence on assemblage structure. The first axis explained most of the species-environment relationship, all variation was explained by the first three canonical axes, and the total intertia of eigenvalues was 0.814 (Table 5). Monte Carlo simulation (1000 iterations) of species-environ- ment correlations were significant for Axis 1 (P = 0.001) and for all canonical axes (P = 0.001). Figure 2 is a CCA biplot where species relationships and environmental factors that were most influential to Page 64 Table 4 Inter-set correlations for 22 environmental variables examined during CCA. Variables in bold were statistically significant (P < 0.05) and used in final analysis for the biplot in Figure 2. Abbreviations used in Figure 2 shown in parentheses. Environmental Variable Axis | Axis 2. Axis 3 Water Depth —0.094 —0.0036 0.0704 Water Velocity (WV) —0.5619 —0.0036 0.0704 Wetted (Spring Brook) Width 0.302 —0.0772 0.2246 (WW) Presence of Palm Roots —0.033 — .01297 —0.0597 Presence of Fines (FINES) 0.5465 0.0196 0.0459 Presence of Sand —0.1301 0.042 —0.1521 Presence of Gravel (GR) —0.3831 0.128 —0.041 Presence of Cobble —0.071 —0.0007 0.1061 CPOM (CPOM) 0.2866 —0.1245 —0.0369 Dissolved Oxygen 0.0511 "0855 —0.3521 Concentration PH 0.1421 —0.1228 0.1427 Water Temperature (TEMP) Electrical Conductance —0.6796 —0.3401 0.0182 —0.0643 —0.1221 0.1527 Bank Angle 0.0965 0.0296 0.1444 Presence of Bank Overhang —0.1707 0.2148 —0.032 Bank Armor Presence (BA) 0.0965 0.0296 0.1444 Grassy Bank Presence 0.0582 —0.2636 0.188 Presence of Other Perennial 0.0596 —0.234 —0.0447 Vegetation on Bank Incised Bank Presence (IB) —0.2142 —0.1887 0.0907 Stable Banks —0.1032. —0.2497 —0.1421 Riparian Cover —0.0439 0.0816 0.0804 Quadrat Position (Bank or 0.2823 —0.2859 —0.0651 Center) (POS) assemblage structure are illustrated by vector length and the association of each species with each environ- mental variable. Vector length illustrates that water velocity, temperature, and the presence of fines and Table 5 Eigenvalues, species-environment correlations, and cumulative percentage variance of species data and species-environment relationship explained by the first three ordination axes of the CCA analysis of the Warm Springs mollusk assemblage. Total Axis I Axis II Axis III Inertia Eigenvalues 0.361 0.123 0.023 0.841 Species-Environment 0.835 0.748 0.471 Correlation Cumulative Percentage Variance Species Data 42.9 57.5 60.3 Species-Environment 71.2 9555 100 Relation The Veligers- Vols 50) Now gravel were the most influential factors. Quadrat position and the presence of incised banks were moderately influential, and wetted width, and the presence of CPOM and armored banks were least important. The plot indicates these species occupied different habitats. Pyrgulopsis avernalis (PA) was associated with higher current velocities (WV), gravel substrate (GR), and higher water temperature (TEMP), and P. carinifera (PC) with moderate current velocities and incised (IB), unarmored banks. Tryonia clathrata (TC) was associated with moderate temperatures, slower currents, bank quadrats, CPOM, and the absence of gravel and fine substrates. Melanoides tuberculata was associated with fine substrate (FINES), wider brooks (WW), cooler temperatures, and lower current velocities. Microhabitat Use and Resource Partitioning: Habitat preference calculations confirm CCA results and indicate these species partition habitat by temperature, water velocity, and substrate. Niche breadth and overlap values suggested that there were differences in diversity of habitat used by each species and that there was common use of some habitats. Pyrgulopsis avernalis occupied a wide diversity of depths (B = 5.21), but preferred depths from 30cm to 40 cm (Figure 3A). It avoided shallow (<15 cm) and deeper water (>45 cm). Niche breadth values indicated that it also occupied a wide variety of current velocities (B = 7.69), but it preferred mean water column velocities > 50 cm/sec and strongly preferred velocities approxi- mately 70-110 cm/sec (Figure 4A). It avoided currents < 40 cm/sec and it was most common in mid-channel quadrats where currents were swift and smaller substrates scarce (Table 6). It also preferred gravel, avoided cobbles, and strongly avoided fines, sand, and CPOM (Figure 5A). It occupied the warmest water temperatures in the spring province, preferred temper- atures near 32°C (B = 1.49), and avoided cooler water (Figure 6A). Pyrgulopsis carinifera also occupied a diversity of depths (B = 5.29), but it preferred habitats < 10 cm deep and avoided depths > 30cm (Figure 3B). It occupied slow and fast currents (B = 5.66) but preferred mean water column velocities from 30 to 40 cm/sec (Figure 4B). Like P. avernalis, it occurred in mid-channel quadrats (Table 6), preferred gravel, avoided sand and CPOM, and strongly avoided fines and cobbles (Figure 5B). It also strongly preferred temperatures near 32°C (B = 1.31) and avoided cooler water (Figure 6B). Tryonia clathrata also preferred the warmest waters (Figure 6C), but occupied substantially different mi- crohabitats than either species of Pyrgulopsis. Tryonia clathrata was most common along spring brook banks where it preferred shallow (<5 cm deep), slow moving D. W. Sada, 2007 S ib P. avernalis 2 ro) PROPORTION °o r=) = rN) 5 15 25 35 45 55 65 Water Depth (cm) S ro) P. carinifera PROPORTION ° 5 15 25 35 45 55 65 Water Depth (cm) Page 65 0.6 T. clathrata © rS PROPORTION ° iy 5 15 25 35 45 55 65 Water Depth (cm) 0.3 M. tuberculata PROPORTION 5 15 25 35 45 55 65 Water Depth (cm) Figure 3. The depth of water occupied by P. avernalis, P. carinifera, T. clathrata, and M. tuberculata. + and ++ = moderately preferred resource categories, respectively, - and — = moderately and strongly avoided resource categories, respectively. B = niche breadth. (<20 cm/sec) water while avoiding deeper, swiftly flowing waters (Figures 3C and 4C, Table 6). Small niche breadth values for depth and velocity (B = 3.50 and 2.82, respectively) indicate its comparatively restricted use of shallow depths and slow currents. It strongly preferred sand, preferred fines and CPOM, and strongly avoided gravel and cobbles (Figure SC). Melanoides tuberculata occupied a wider diversity of water temperatures than the springsnails (B = 2.85), and it strongly preferred temperatures near 25°C and strongly avoided the warm temperatures preferred by springsnails (Figure 6D). Differences between springs- nail and M. tuberculata preferences for temperature probably account for the importance of temperature in structuring the assemblage that was shown by CCA (Figure 2). It exhibited no preference for water depth but it strongly avoided habitats deeper than 30 cm (Figure 3C). It was most common along springbrook banks (Table 6). Current velocities from 0—10 cm/sec were preferred and mean water column velocities < 10 cm/sec were strongly preferred (Figure 4D). It showed no preference or avoidance of CPOM. Melanoides tuberculata used habitats similar to those occupied by T. clathrata and quite different from both Pyrgulopsis_species. Although habitats used by T. clathrata and M. tuberculata were similar, M. tubercu- lata strongly preferred the presence of fine substrate and strongly avoided sand, gravel, and cobble (Fig- ure 5D). These two species also appeared to occupy habitats. with different temperatures and current velocities (Figures 3D and 4D, respectively). Tryonia clathrata occupied shallow habitats (Figure 2D) where sand substrate was associated with slightly swifter current while M. tuberculata occupied habitats where fine substrates were associated with very low current velocities. These characteristics suggest that M. tuber- culata may be relatively tolerant of nocturnal decreases in dissolved oxygen concentrations that can be associated with fine substrates. Use of this habitat type is consistent with observations of its habitat use by Dudgeon (1989), Gutiérrez et al. (1997), and Duggan (2002). Differences in habitat use by springsnails and M. tuberculata suggest that their interactions may be minimal at Warm Springs. Niche overlap values generally confirm habitat use shown in Figures 3—S. Values were < 0.5 between T. clathrata and both species of Pyrgulopsis for water depth and substrate type (with exception of 0.638 between T. clathrata and P. carinifera for depth), which suggests that intra-generic habitat use of species in this Page 66 0.3 P. avernalis + 2 iy PROPORTION oO 0 20 40 _ 60 80 100 Water Velocity (cm/sec) 0.3 P. carinifera S iy PROPORTION ° 0 20 40 60 80 100 Water Velocity (cm/sec) The Veliger, Vol. 50, No. 2 0.5 > 0.4 2 T. clathrata fr 0.3 & o 2-2 oc OA 0 0 20 40 60 80 100 Water Velocity (cm/sec) 0.5 > 0.4 2 T. clathrata = 0.3 m4 © 0-2 a eon 0 0 20 40 60 80 100 Water Velocity (cm/sec) Figure 4. Mean water column velocity of habitats occupied P. avernalis, P. carinifera, T. clathrata, and M. tuberculata. Preference and avoidance illustrated as in Figure 3. springsnail assemblage is minimal (Tables 7 and 8). Niche overlap values for P. avernalis and P. clathrata were low for current velocity (<0.35) and > 0.5 for water depth and substrate type, which indicates that habitat use by these species is similar and differs primarily in the use of fast and moderate currents by P. avernalis and P. clathrata, respectively. Overlap be- tween the springsnail assemblage and M. tuberculata was high for water depth, but overlap was also high between velocities used by the melania, P. carinifera, and T. clathrata (Table 7). Highest overlap in current velocity occurred between T. clathrata and M. tubercu- lata, and lowest was between P. avernalis and M. tuberculata (Table 7). Overlap for fines, gravel, and cobble substrate use was relatively high among all species in the assemblage, but overlap was low between both Pyrgulopsis species and T. clathrata for use of sand (Table 8). Overlap for use of sand was also low between M. tuberculata and T. clathrata (Table 8), which confirms results of calculations showing M. tuberculata and T. clathrata preferred fines and sand, respectively. DISCUSSION Springsnails represent the most diverse family of gastropods in western North America and many species occupy the smallest aquatic habitats in the most arid regions of the continent. In these areas, most populations inhabit isolated springs and spring prov- inces on valley floors and along the base of mountain blocks. Populations rarely inhabit springs higher than 2,400 m elevation. Temperature and EC of springsnail habitats range from 10°C to 40°C and from 70 umhos/ cm to 37,000 umhos/cm, respectively (Hershler and Sada, 1987; Sada and Deacon, 1995; Hershler, 1998). The amount of habitat occupied by _ springsnail populations ranges from < 1 m? in small springs to > 100 m? in large springs. Hershler (1998) estimated that the density of some populations may reach 10,000/m°. Most populations are relictual in aquatic systems that have persisted since ancient pluvial periods, and they are restricted to springs with minimal environmental Table 6 Proportion of individuals of each species of mollusk occurring in spring brook mid-channel and bank quadrats sampled at Warm Springs. Species Mid-Cannel Bank P. avernalis 0.80 0.20 P. carinifera 0.53 0.47 T. clathrata 0.18 0.82 M. tuberculata 0.24 0.76 D. W. Sada, 2007 Page 67 P. avernalis - ‘= PROPORTION FINES SAND GRAVEL COBBLE CPOM Substrate Types 1 P. carinifera 0.8 a 2) EF 0.6 (om m4 ro) 0.4 co a 0.2 0 FINES SAND GRAVEL COBBLE CPOM Substrate Types T. clathrata 2a OQ = oa fe) a re) oc a FINES SAND GRAVEL COBBLE CPOM Substrate Types 1 M. tuberculata 0.8 =a 2) 0.6 fam 4 00.4 ao a 0.2 FINES SAND GRAVEL COBBLE CPOM Substrate Types Figure 5. The proportion of P. avernalis, P. carinifera, T. clathrata, and M. tuberculata that were associated with the presence (L)) and absence (™) of different substrates. Preference and avoidance shown only for occupation of quadrats with the presence of a substrate type listed. Preference and avoidance illustrated as in Figure 3. variability (Taylor, 1985). As with other crenobiontic species, densities are greatest near spring sources where physicochemical environments are relatively stable compared to downstream reaches of spring brooks where seasonal and daily environmental variability is relatively high and springsnails are sparse or absent (Noel, 1954; Hershler, 1998; McCabe, 1998). They do not inhabit reaches that dry on a regular basis, but they may colonize downstream reaches where stressful events such as flooding and drying are infrequent. Past quantitative studies suggest that springsnails preferentially occupy limited portions of springs with suitable chemistry. In laboratory experiments O’Brien and Blinn (1999) demonstrated that P. montezumensis inhabited a limited reach of spring brook where CO; concentrations ranged from 110-315 mg/L. They did not occupy upstream reaches where concentrations were greater and downstream reaches where concen- trations were less. Annual patterns of variability in P. bruneauensis density and its preference for upstream portions of spring brook were attributed to water temperature- by Mladenka and Minshall (2001). In experiments with springsnail assemblages in mound springs of Australia, Ponder et al. (1989) demonstrated the influence of several environmental factors on activity levels and survivorship of one amphibious species and several aquatic species. Each of these species exhibited intolerance to desiccation, low and elevated salinity concentrations, deoxygenated water, low and elevated water temperature, and to varying exposure to submersion. Behavioral responses to light also varied among species. These studies also recorded the relative abundance of several species in eight zones (e.g., spring source, upper part of outflow, middle part of outflow, etc.) in a number of springs. Patterns of zonal occupation were weak, but each assemblage typically included one amphibious species and one large species, and from one to three small aquatic species. They also concluded that springsnail niche potential was fully exploited in these springs and that introduction of species from other springs was therefore unlikely. The study at Warm Springs showed that structure of this assemblage of native and non-native mollusks was influenced by water temperature and several physical Page 68 The Veliger, Vol. 50, No. 2 0.9 2 ® P. avernalis PROPORTION 2 w 24 25 26 27 28 29 30 31 32 Water Temperature (°C) S & P. carinifera S 2) PROPORTION Oo az So iy 24 25 26 27 28 29 30 31 32 Water Temperature (°C) 0.9 2 fo2) T. clathrata PROPORTION 0.3 24 25 26 27 28 29 30 31 32 Water Temperature (°C) M. tuberculata PROPORTION 24 25 26 27 28 29 30 31 32 Water Temperature (°C) Figure 6. Water temperature of habitats occupied by P. avernalis, P. carinifera, T. clathrata, and M. tuberculata. Preference and avoidance illustrated as in Figure 3. elements of the aquatic and spring brook bank environment. These conclusions appear to confirm qualitative observations made during taxonomic stud- ies. They are also consistent with observations by Mladenka and Minshall (2001) that water temperature was an important element of their occupied habitat. Although springsnails at Warm Springs preferred warm water, preferences for temperature did not differ among these species and temperature was not an important factor segregating springsnail habitat use. Springsnails at Warm Springs occupied a wide diversity of aquatic habitats, which was indicated by relatively high niche breadth values for each species in several habitat elements. Depths, velocities, and sub- strates occupied by 7. clathrata were generally more specific than those occupied by either species of Pyrgulopsis. Both Pyrgulopsis generally occupied deep- er and swifter water, and larger substrates than 7. clathrata, which preferred water < 5 cm deep, velocities < 20 cm/sec, and sand and fine substrates. Additional partitioning among assemblage members is suggested by the use of mid-channel and springbrook margin habitats. Although each species occurred across springbrooks, most T. clathrata occurred along spring brook margins, P. carinifera was relatively evenly distributed between margins and the mid-channel, and P. avernalis was most abundant in mid-channel habitats. This appears to be consistent with habitat Table 7 Niche overlap values for water depth and mean water column velocity among mollusks in Warm Springs, Clark County, Nevada. Abbreviations for species as shown in Figure 2. Water Depth PA PC TC MT PA .678 .469 649 PC 366 .638 .789 TC BLOF 482 =f9) MT 159 .502 877 Mean Water Column Velocity D. W. Sada, 2007 Table 8 Niche overlap values for fines, sand, gravel, and cobble substrates among mollusks in Warm Springs, Clark County, Nevada. Abbreviations for species as shown in Figure 2. Fines/Sand PA PC TC MT PA .961/.979 .635/.103 .404/.994 PC .818/1 .674/.124 .443/.973 TC .510/1 .692/1 .769/.097 MT .500/.991 .500/.991 .500/.991 Gravel/Cobble preferences exhibited by each species. Tryonia clathrata occupied shallow, slow habitats and fine substrates that occured along springbrook banks. Pyrgulopsis avernalis was most abundant in mid-channel habitats where substrates are larger and depths and current velocities greatest and Pyrgulopsis carinifera preferred slower water and moderate depths that are lateral habitats in mid-channel and along springbrook margins. Prefer- ential occupation of margin and mid-channel habitats suggests that these springsnails may exhibit zonal preferences of habitat use. Differences between this observation and conclusions by Ponder et al. (1989) that zonal preferences were not apparent may be attributed to sample techniques. The wide range in springsnail density within 120 cm? quadrats (range from 0 to 346 springsnails) at Warm Springs suggests there is wide spatial variability in springsnail abun- dance in a spring and that sample methods using large quadrats may yield weak relationships between springs- nail abundance and characteristics of the spring environment. Determining these relationships appears to require quantitative sampling within a small area. Sampling springsnail abundance and habitat charac- teristics within 120 cm? quadrats appears suitable to examination these relationships. Observations at Warm Springs provide insight into the potential effects of the M. tuberculata on springs- nail populations. Observations of potential competitive interactions between M. tuberculata and several species of Tryonia and Pyrgulopsis in Ash Meadows, another thermal spring province in southern Nevada, that were hypothesized by Hershler and Sada (1987) were not confirmed at Warm Springs. Niche overlap between MM. tuberculata and both species of Warm Springs Pyrgu- lopsis was small for all measured habitat elements. Overlap between M. tuberculata and T. clathrata was more extensive with both species occupying marginal, slow moving habitats with small substrate. In spite of these similarities, interactions appeared to be minor because they utilized different temperatures, substrates, and water velocities. Melanoides tuberculata preferred Page 69 cooler water, finer substrate, and slower currents than T. clathrata. Differences in habitat use among springs- nails and M. tuberculata suggest that competitive interactions between these mollusks are relatively minor and that presence of the M. tuberculata minimally influences the abundance or habitat use of springsnails at Warm Springs. Physiological requirements of springsnails demon- strated by Ponder et al. (1989), O’Brian and Blinn (1999), and Mlandeka and Minshall (2001) and springsnail preference for physical components of the environment at Warm Springs suggest that each taxon may be restricted to portions of a spring that provide suitable physicochemical conditions. Additionally, it suggests that springsnail abundance and distribution may be a function of factors that alter these conditions. These observations have wide implications for springs- nail biogeography and conservation by suggesting that each taxon may be adapted to comparatively specific physicochemical aspects of their ‘home’ springs. These rather specific adaptations may be important factors limiting springsnail dispersal and restricting most taxa to springs with similar environments within a limited geographic area. Several recent articles have noted the vulnerability of springsnails to extirpation because of their limited distribution and life history requirements (e.g., Hershler, 1998; Hurt and Hedrick, 2004). Studies at Warm Springs provide quantitative evidence that springsnail abundance may be affected by any factor affecting water temperature (e.g., springbrook diver- sion, integrity of riparian vegetation), and the quality and heterogeneity of spring habitats. Human activities that reduce environmental heterogeneity (e.g., reduce discharge, channelize, or alter springbrook bank morphology and vegetation) are likely to reduce springsnail abundance or extirpate populations because they alter elements of the environment that define springsnail habitat. Effects of reduced habitat quality and heterogeneity by channelization, siltation, and diversion on springsnail abundance are apparent at Warm Springs where springsnails are scarce or absent from approximately 85 percent of historically occupied springbrooks. In spite of these declines, these springs- nail populations appear to be comparatively resilient because their abundance rapidly increased following springbrook restoration for Moapa dace. Reestablish- ing springsnails in their historic range will require restoring springbrook characteristics and minimizing factors that reduce environmental heterogeneity, such as decreased discharge attributed to diversion and groundwater use. Over the past decade, Scoppettone (1993) and Scoppettone et al. (1992) delineated characteristics of Moapa dace habitat use, and showed that adults are most common in deep, comparatively large habitats where they can hold near mid-water column and feed Page 70 on drifting macroinvertebrates. This information has been integrated into conservation programs to enhance and protect Moapa dace from non-native fishes and activities that have adversely modified springbrooks and the Muddy River. Springsnail preference for relatively shallow habitats with diverse substrate composition suggests that springbrook restoration designed solely for Moapa dace may not provide sufficient heterogeneity for springsnails. The declining status of springsnails and_ their sensitivity to habitat alteration is an indicator of the ecological consequences of activities that degrade springs. This suggests that changes in management are necessary to maintain biotic integrity and prevent future declines in crenobiontic species and habitats that support a large portion of the aquatic biodiversity in arid lands. Acknowledgments. This study was funded by The Nature Conservancy, Southern Nevada Projects Office as a compo- nent of programs to conserve arid land spring systems. D. Herbst assisted with fieldwork and L. Sada assisted with laboratory identification. The manuscript was improved by comments from J. Furnish, S. Sharpe and C. Rosamond. LITERATURE CITED Burcu, J. B. & J. L. TOTTEHAM. 1980. 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This paper presents a review of the taxonomy of the species belonging to the genus Cerithiopsis Forbes & Hanley, 1850 from Brazil, that are characterized by the two adapical rows of nodules in each teleoconch whorl fused together on the initial teleoconch whorls, then becoming gradually separated on the subsequent ones. The following species are reported for the first time from Brazilian coast: C. fusiformis (C. B. Adams, 1850), originally described from Jamaica; and C. aimen Rolan & Espinosa, 1995 and C. prieguei Rolan & Espinosa, 1995, both originally from Cuba. Two new species are described: Cerithiopsis balaustium and Cerithiopsis capixaba, both from the northeast-southeast coast of Brazil. The five species have minor differences in shell shape and sculpture pattern, being easily distinguished by the protoconch sculpture: in C. fusiformis it is smooth, with a thin spiral keel in the middle on two abapical whorls; C. aimen has a similar smooth protoconch, but lacking any spiral keel; C. prieguei bears two thin spiral cords on adapical whorls, connected by thin axial riblets, also, the sutural area of the protoconch is microscopically granulated; C. balaustium has a small protoconch with small axial riblets that do not touch the suture; C. capixaba has an elongate protoconch with initial whorls finely granulated and subsequent ones with axial riblets that connect the suture. Key Words: Cerithiopsidae, Cerithiopsis, Brazil, South America, taxonomy. INTRODUCTION The genus Cerithiopsis Forbes & Hanley, 1850 is contains marine microgastropods belonging to the family Cerithiopsidae H. & A. Adams, 1853. Along with the family Triphoridae Gray, 1847, Cerithiopsidae composes the largest part of the superfamily Triphor- oidea Gray, 1847 (Ponder & Warén, 1988; Ponder, 1998). The characteristics used in the taxonomy and identification of species of Cerithiopsis, particularly the sculpture pattern, are not easily observed, and worn shells or shells lacking the protoconch are often impossible to identify (Laseron, 1951; Marshall, 1978). There is no consensus concerning the supraspecific classification of Cerithiopsis. Some authors (e.g., Jay & Drivas, 2002, following Marshall, 1978) restricted the concept of the genus, adopting other generic names, such as Joculator Hedley, 1909, Horologica Laseron, 1956, Mendax Finlay, 1927 and Prolixodens Marshall, 1978: Odé (1989) also considered Joculator, but at the subgeneric level. Herein, we adopted Cerithiopsis in a broad sense, following Rolan & Espinosa (1995) and Rolan et al. (2007). We avoided the use of other generic names, because we did not have access to soft parts or radulae, the analysis of which seems to be essential for proper supraspecific classification in this group. Rolan & Espinosa (1995) and Rolan et al. (2007) distin- guished, for working purposes, “groups of species” based on color patterns (e.g., brown colour species, banded and variable colored species). Whereas most species of this genus have three rows of nodules, equidistant or almost equidistant along the entire teleoconch, some species have the two adapical rows of nodules fused together on the first whorls and becoming gradually separated on the subsequent ones, as described for Cerithiopsis fusiformis (C. B. Adams, 1850), a widely distributed species in the western Atlantic. During studies on the taxonomy of Brazilian Cerithiopsidae, we found some shells that exhibit such characteristcs of teleoconch sculpture, but can be distinguished by other traits. This paper presents the description of two new species from Brazil, as well as the first, local occurrence of three other species that were originally described from the Caribbean region. MATERIAL AND METHODS The material used for this paper was collected in several localities along the Brazilian coastline, and is listed R. Figueira & A. Pimenta, 2007 separately for each species, with the number of shells in brackets. The study was based entirely on conchologi- cal analyses. The terminology and characters used to identify the species were based on Laseron (1951, 1956), Marshall (1978), Rolan & Espinosa (1995) and Jay & Drivas (2002). Abbreviations used: MCZ: Museum of Comparative Zoology, Cambridge; MNCN: Museo Nacional de Ciencias Naturales, Madrid; IBUFRJ: Instituto de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro; MNHN: Muséum National d Histoire Naturelle, Paris; MNRJ: Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro; MORG: Museu Oceanografico ““Eliézer de Carvalho Rios” da Funda¢ao Oceanografica do Rio Grande, Rio Grande; MZSP: Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo; REVIZEE: Recursos Vivos da Zona Economica Exclusiva; sta: collection station number; NOAN: Navio Oceanografico ‘Antares; NOWB: Navio Oceanografico ““Professor Wladimir Besnard.” SYSTEMATICS Family Cerithiopsidae H. & A. Adams, 1853 Subfamily Cerithiopsinae H. & A. Adams, 1853 Genus Cerithiopsis Forbes & Hanley, 1850 Cerithiopsis Forbes & Hanley, 1850. Type-species by original designation: Cerithium tubercularis Montagu, 1803; Recent, Europe. Diagnosis: Shell of various shapes (conic to ellipsoidal), protoconchs smooth or sculptured with axial and/or spiral ridges, or finely granulated; teleoconch whorls sculptured by usually three spiral cords, crossed by axial ribs, forming small nodules, with various degrees of conpicuousness. Aperture subquadrangular to cir- cular; siphonal canal short and oblique. Cerithiopsis fusiformis (C. B. Adams, 1850) (Figures 1—5) Cerithium fusiforme C. B. Adams, 1850:120—-121; Clench & Turner (1950:285, pl. 38, fig. 4). Cerithiopsis fusiforme: Usticke (1959:42, not illustrat- ed); Jong & Coomans (1988:46, not illustrated); Rolan & Espinosa (1995:132, figs. 2-6, 47). Cerithiopsis (Cerithiopsis) fusiformis: Vokes & Vokes (1983:18, pl. 27, fig. 6). Joculator fusiformis: Redfern (2001:74, pl. figs. 310A— B). Types: Holotype: MCZ 186127, Caribbean Sea, Ja- maica; C.B. Adams coll. Page 73 Material examined: The holotype and: —Pernambuco: MNHN, [2] Enseada dos Corais, 1984-1989, P. Maestrati col.; —Bahia: IBUFRJ 12906, [2] Baia de Todos os Santos, sta 56vv, 12/iv/1997; IBUFRJ 12907, [1] Baia de Todos os Santos, sta Ilvv, 22/iv/1997; IBUFRJ 12908, [18] Baia de Todos os Santos, sta 54vv, 29/V/1997; IBUFRJ 12909, [9] Baia de Todos os Santos, sta S55vv, 03/iv/1997; IBUFRJ 13887, [1] Baia de Todos os Santos, 5/iv/1997; MORG 48812, [3] R. Areia, Abrolhos, 1/1985, Eq. MORG coll.; MORG 48811, [1] I. Guarita, Abrolhos, 11/1987, A. Silveira and L. Laurino coll.; — Espirito Santo: IBUFRJ 12905, [1] REVIZEE sta c65 (18°53'37’S, 39°06'23”W, 50 m), 25/ iv/1996, NOAN coll.; IBUFRJ 9515, [1] REVIZEE V sta c64 (19°17'42”S, 38°02'06"W, 63 m), 25/iv/1996, NOAN coll.; IBUFRJ 12903, [12] REVIZEE sta vv38 (19°44’S, 38°24’W, 71.4 m), 29/11/1996, NOAN coll; MNRJ 11887, [3] REVIZEE sta vv38 (19°44’S, 38°24'W, 71.4 m), 29/11/1996, NOAN coll.; MORG 48816, [39] Aracruz, 1/viii/1988, Vera Abud coll.; IBUFRJ 12904, [6] REVIZEE sta c62 (20°30'02’S, 37°28'51”W, 96 m), 25/iv/1996, NOAN coll.; IBUFRJ 10049, [1] REVIZEE V_ sta c61 (20°30'38’S, 37°19'06"W, 88 m), 24/iv/1996, NOAN coll.; IBUFRJ 12900, [1] REVIZEE sta vv21 (20°38’S, 40°00'W, 33m), 26/11/1996, NOAN coll.; —Rio de Janeiro: IBUFRJ 13665, [3] Praia da Figueira, Angra dos Reis, 1998, C. Alvarenga coll.; MORG 41970, [1] off Macaé, sta 3207, 12/1v/1997, Oceanographic Vessel “Victor Hensen” coll. Cerithiopsis aimen Rolan & Espinosa, 1995 (Figures 6—11) Cerithiopsis aimen Rolan & Espinosa, 1995:142, figs. 27-30; Boyko & Cordeiro (2001:27). Types: Holotype: MNCN_ 15.05/17220, Cienfuegos Bay, Cuba; one paratype at AMNH 226504. Material examined: The holotype and: IBUFRJ 12902, [2] REVIZEE sta vv31 (18°52'S, 39°35'W, 22.8 m), 28/ 11/1996, NOAN coll.; MNRJ 11889, [1] REVIZEE sta vv31 (18°52’S, 39°35'W, 22.8 m), 28/11/1996, NOAN coll.; IBUFRJ 15373, [1] REVIZEE VI sta y7 (77 m): IBUFRJ 15565, [1] Baia de Todos os Santos, Bahia state, S/iv/1997; IBUFRJ 15564, [4] Baia de Todos os Santos, Bahia state sta Il vv, 22/iv/1997. Cerithiopsis prieguei Rolan & Espinosa, 1995 (Figures 12-18) Cerithiopsis prieguei Rolan & Espinosa, 1995:142, figs. 31-34, 45; Boyko & Cordeiro (2001:28). Page 74 The Veliger, Vol. 50, No. N Figures 1-5. Cerithiopsis fusiformis. 1. holotype (MCZ 186127); 2-3, 5. IBUFRJ 12906; 4. IBUFRJ 12907; 1—2. whole shells (iengihs: 3.0 mm); 3-4. protoconchs; 5. last whorl. Scale bars: 200 um. Fis »6-11. Cerithiopsis aimen. 6-7. holotype (MNCN 15.05/17220); 8-11. IBUFRJ 12907; 6, 8. whole shells (respective lengths: 3.2 i, 2.1 mm); 7, 10-11. protoconchs; 9. last whorl. Scale bars: 200 tum. R. Figueira & A. Pimenta, 2007 Page 75 Figures 12-18. Cerithiopsis prieguei. 12, 16. holotype (MNCN 15.05/17221); 13-15, 17-18. IBUFRJ 14139; 12-14. whole shells (respective lengths: 2.2 mm, 2.5 mm, 2.0 mm); 15. last whorl; 16-17. protoconchs; 18. detail of early protoconch whorls. Scale bars: 15-17: 200 um; 18: 50 um. Types: Holotype: MNCN 15.05/17221, La Habana, Cuba; several paratypes listed in Rolan & Espinosa (1995). Material examined: The holotype and: IBUFRJ 14139, [3] REVIZEE sta vv38 (19°44'S, 38°22'W, 71.4 m), off Espirito Santo state, 29/11/1996, NOAN col.; MNRJ 11888, [3] REVIZEE sta vv38 (19°44’S, 38°22’W, 71.4 m), off Espirito Santo state, 29/11/1996, NOAN col. Cerithiopsis balaustium n.sp. (Figures 19—23) Type material: holotype: MZSP 86307 REVIZEE sta 6666 (24°17.129'S, 44°12.149'W, 163 m), off Sao Paulo state; paratypes: IBUFRJ 15223, [1] Baia de Todos os Santos, Bahia state sta Ilvv, 22/iv/1997; IBUFRJ 15566, [1] Baia de Todos os Santos, Bahia state, sta 54vv, 29/v/1997; IBUFRJ 15371, [1] REVIZEE VI sta y7 (77m); MZSP 86306 [3] REVIZEE sta 6666 (24°17.129'S, 44°12.149'W, 163m), off Sao Paulo state; MNRJ 10981, [1] PADCT sta 6577 (25°15.76’S, 45°04.62'W, 124 m), off Sao Paulo state; MZSP 86308 [1], PADCT sta 6595 (26°23’S, 46°39'W, 175 m), off Santa Catarina state. Type locality: off Sao Paulo state, southeast coast of Brazil (24°17.129’S, 44°12.149’W, 163 m). Etymology: From Latin Balaustium = balauster. The species is named after its protoconch sculpture, whose series of axial riblets resembles the architectural elements of a balustrade. Diagnosis: protoconch with small axial riblets restricted to the middle portion of each whorl, not touching the suture; first teleoconch whorls with the two adapical rows of nodules fused together and becoming separated on the subsequent whorls. Description: shell small, reaching 2mm of height, somewhat pupiform, opaque. Protoconch subcylindri- Page 76 The Veliger, Vol. 50, No. 2 Figures 19-23. Cerithiopsis balaustium n.sp. 19, 21-23. holotype (MZSP 86307); 20. paratype (IBUFRJ 15223); 19-20. whole shells (respective lengths: 1.3 mm, 1.4 mm); 21. last whorl; 22. protoconch; 23. detail of protoconch sutural area. Scale bars: 21—22: 200 um; 23: 50 um. Figures 24-28. Cerithiopsis capixaba n.sp. 24-25, 28. holotype (IBUFRJ 15221); 26-27. paratype (MZSP 86309); 24. whole shell (length: 2.5 mm); 25-26. protoconchs; 27. detail of early protoconch whorls; 28. last whorl. Scale bar: 25-27: 200 um; 28: 100 um. R. Figueira & A. Pimenta, 2007 cal, dark yellow, with about 3.5 whorls of convex outline, the first one dome-shaped; two abapical whorls with small prosocline axial riblets which are restricted to the middle area of the whorls, without reaching the suture, except in its last half-whorl; sutural area with granules, especially on its adapical region, which become thicker and coalescent on the third whorl, forming a small spiral suprasutural cord. Teleoconch with up to five whorls; color varying from light to dark yellow. Suture well impressed. Teleoconch whorl sculpture formed by three spiral cords and about 16 axial ribs on the fourth whorl with the formation of rounded nodules on the intersections; early whorls with the two posterior cords fused together, their nodules appearing to be a single row of bilobate ones, these cords gradually separate from each other in the subsequent whorls until they are equidistant on the last one. Base short, with a slightly nodulose spiral cord on its periphery and two large spiral grooves, separated by a spiral cord, axial growth lines, and thin spiral lines in its anteriormost portion. Aperture somewhat ellip- tical, with a short siphonal canal. Outer lip thickened. Cerithiopsis capixaba n.sp (Figures 24-28) Type material: holotype: IBUFRJ 15221, [1] REVIZEE sta vv21 (20°38’S, 40°00’W, 33 m), off Espirito Santo state, 26/1/1996, NOAN coll.; paratypes: IBUFRJ 9275, [1] REVIZEE sta c63 (19°40'42”S, 38°08'15’W, 61m), off Espirito Santo state, 25/iv/1996, NOAN coll.; IBUFRJ 14138, [2] REVIZEE sta vv38 (19°44’S, 38°22'W, 71.4 m), off Espirito Santo state, 29/11/1996, NOAN coll.; MNRJ 10982, [3] REVIZEE VI sta y7 (77 m); MZSP 86309, [1] REVIZEE sta 6662 (24°00.946'S, 43°55.540’W, 135m), off Sao Paulo state. Type locality: off Espirito Santo state, southeast coast of Brazil (20°38’S, 40°00’ W, 33 m). Etymology: ““Capixaba”’ is the common denomination given to natives of the state of Espirito Santo, in southeastern Brazil. Diagnosis: protoconch with many thin axial ribs covering the entire surface of each whorl; early teleoconch whorls with the two adapical rows of nodules fused together and becoming separated on the subsequent whorls. Description: shell small, reaching 3 mm of height, pupiform with a somewhat acuminate apex, opaque. Protoconch cylindrical, yellow, with about 5 whorls of convex outline, the first one dome-shaped; first two whorls with small granules organized in spiral rows, subsequent_whorls with prosocline axial riblets reach- Page 77 ing the suture. Teleoconch with up to seven whorls; color light caramel. Suture well impressed. Teleoconch whorl sculpture formed by three spiral cords and about 16 axial ribs on the sixth whorl with the formation of rounded nodules on the intersections; early whorls with the two posterior cords fused together, their nodules appearing to be a single row of bilobate ones, these cords gradually separate from each other in the subsequent whorls until they are equidistant on the sixth whorl. Base very short, with a spiral cord on its periphery and two large spiral grooves, separated by a spiral cord; axial growth lines, and spiral lines in its anteriormost portion. Aperture subquadrate, with a short siphonal canal. Outer lip thickened. DISCUSSION Marshall (1978), studying the Cerithiopsidae from New Zealand, considered several genera apart from Cer- ithiopsis, such as Joculator, Horologica, Prolixodens, among others. The distinction among these generic names was based on a combination of teleoconch shape and sculpture, as well as on protoconch type. In a similar way, Jay & Drivas (2002) considered several generic names, but relied mainly on protoconch characteristics to restrict the concept of Cerithiopsis to shells with smooth or punctated protoconchs, along with the genera Joculator and Horologica. The distinc- tions among these three genera were related to the general shape of the shell and the base, and the number of spiral nodulose rows per whorl. Species with sculptured protoconchs, on the other hand, were assigned to the genera Dizoniopsis Sacco, 1895, Mendax and Prolixodens. However, the concept adopted for some genera, such as Mendax, are not in accordance with those used by Marshall (1978), who regarded Mendax as species with lecithotrophic larval type of few (2 4) whorls, with non-granulated earlier whorls. The species included by Jay & Drivas (2002) have 3—4 whorls (planktotrophic type), with granulated earlier whorls. Furthermore, Marshall (1978) stated that classifica- tions based on protoconch types do not reflect phylogeny, since there can be genera with both kinds of development (1.e., planktotrophic and _lecitho- trophic), reflecting different protoconch sculpture patterns. In the absence of radular and/or anatomical data, we used Cerithiopsis in a wide sense to encompass both smooth or axially sculptured protoconchs, a position adopted by Rolan & Espinosa (1995), Also, we recognized no distinction between Cerithiopsis and Joculator, because the differences among them are related to general shell shape, which is quite variable in some species of Cerithiopsis. Cerithiopsis s.1. species are generally characterized by teleoconch sculpture of three or two rows of nodulose Page 78 The Veliger, Vol. 50, No. 2 cords, nearly equidistant, on each whorl. In some species, however, the two adapical nodulose rows are very close to each other, and quite distant from the abapical row; in some cases, these two adapical rows may be fused in a kind of double row, or even only one row may be visible, especially on earlier teleoconch whorls. Marshall (1978) included species with this teleoconch sculpture in Horologica, defined as tele- oconchs with two spiral cords, a third emerging by fission and subsequent development of the first spiral cord. However, Marshall (1978) himself expressed doubt as to whether Horologica would prove to be a subgenus of Joculator after eventual forthcoming anatomical data. This suggests to us that the number and disposition of spiral cords are also insufficient to discriminate among generic entities, and again, we have considered Cerithiopsis in a broad sense. Typical western Atlantic representatives of the type of sculpture described above, are Cerithiopsis fusiformis (C. B. Adams, 1850) (Figures 1—5), from Jamaica; and Cerithiopsis aimen (Figures 6-11) and = Cerithiopsis prieguei (Figures 12-18), both from Cuba, described by Rolan & Espinosa (1995). These three species are herein reported for the first time from the Brazilian coast. Rolan & Espinosa (1995) stated that the protoconch of C. fusiformis seems smooth, but has a_ spiral angulation in its middle and, in some specimens, an additional spiral cord near the suture. Some level of erosion may be responsible for this variation. These protoconch characters are visible in the Brazilian specimens (Figures 3, 4). Cerithiopsis aimen (Figures 6-11) can be distin- guished by its smooth protoconch, which lacks any spiral sculpture (Figures 7, 10, 11). In addition, as discussed by Rolan & Espinosa (1995), C. aimen also differs from C. fusiformis in the arrangement and relative size of the nodules on the first teleoconch whorl. The first teleoconch whorl of C. aimen has three, equally spaced, nodulose rows, the middle one smallest, until the second teleoconch whorl, when the two upper rows become fused (Figures 7, 10, 11). In C. fusiformis, the adapical row is smaller than the other two and is already closer to the middle row from the beginning of the teleoconch (Figures 3, 4). Cerithiopsis prieguei (Figures 12-18) displays the same sculptural pattern on the first teleoconch whorl (Figures 16, 17) as C. fusiformis. The easiest way to distinguish this species from the others is by the protoconch sculpture, which has two fine, spiral cords, with small, incomplete, oblique axial ribs (Figures 16, 17). Further, the sutural area of the protoconch has some small granules (Figure 18). Besides the three species mentioned above, this paper presents the description of two new species, with similar shell shape and sculpture pattern. Both Cerithiopsis balaustium and Cerithiopsis capixaba have the adapical row smaller than the other two at the beginning of the teleoconch, similar to C. fusiformis. Particularly in Cerithiopsis balaustium the adapical rows of nodules are extremely close since the first whorl. The teleoconch whorls of the species studied are very similar, though their shapes are not exactly the same: Cerithiopsis balaustium (Figures 19, 20) is somewhat globose and smaller than the others, whereas Cerithiop- sis capixaba (Figure 24) is somewhat pear-shaped, and C. fusiformis (Figures 1, 2), C. aimen (Figures 6, 8) and C. prieguei (Figures 12-14) are oval. Still, in our opinion, this trait is too variable and hardly sufficient to allow a proper identification of these species. There are also slight differences in the separation of the adapical rows of nodules. In C. fusiformis, C. aimen, C. prieguei and C. capixaba, the rows become separated in the fifth or sixth whorls only; whereas in C. balaustium the two fused spiral cords are already separated by the fourth whorl (Figures 19, 20). It is clear, though, that among the conchological characters, the sculpture on the protoconch is the most reliable and conclusive when telling these species apart. All the species have protoconchs of the planktotrophic type, with four or five whorls. Marshall (1978) stated that, although not reliable for generic classification, differences in sculpturing of the planktotrophic proto- conch may be used for species discrimination, because species with this larval development are usually intraspecifically constant. The protoconch of Cerithiopsis balaustium has short axial cords that do not touch the suture (Figures 22, 23), whereas that of Cerithiopsis capixaba has five whorls, the two adapical finely granulose and the remainder with axial cords touching the suture above and below (Figures 25-27). The protoconch sculpture of C. balaustium is somewhat similar to that of Prolixodens sknips Jay & Drivas, 2002, but in this species, which occurs in the Indian Ocean, the axial riblets are more prominent; besides, the teleoconch does not have the two fused adapical rows in the adapical whorls. A recently described species from Cuba, Cerithiopsis apexcostata Rolan, Espinosa & Fernandez-Garceés (2007) has a very similar protoconch sculpture, but in this species, the axial cords, touch the suture below, while in C. balaustium, the cords are, in most of the protoconch whorls restricted to the middle area of the whorl. Also, in C. apexcostata, the two posterior cords in the teleoconch whorls never fuse together, as occurs in the three older whorls of C. balaustium. Also, for C. capixaba, a similar protoconch can be found in three species from the Indian Ocean, Mendax penneyi, Mendax mascarenensis and Mendax ribesae, all described by Jay & Drivas (2002); however, the teleoconch shapes and sculpturing are markedly R. Figueira & A. Pimenta, 2007 Page 79 different. Rolan et al. (2007) described and illustrated the protoconch of Cerithiopsis ara Dall & Bartsch, 1911, with 4 % whorls, the first one with spiral cords and the subsequent with axial ribs crossed by small spiral threads; this is very similar to the protoconch of C. capixaba, but in the species from Brazil, the spiral cords in the initial whorls are formed by small granules organized in spiral rows, and in the subsequent whorls, there are no spiral threads. The most important difference between the two species, however, is in the general shell shape, elongate and with an accuminate apex in C. capixaba and short and pupoid in C. ara. Cerithiopsis prieguei has a protoconch ornamented with small granules at the sutural area of the two earlier whorls and two spiral cords connected by thin prosocline axial riblets on the two abapical whorls (Figures 16-18). The protoconchs of C. fusiformis (Figures 3, 4) and C. aimen (Figures 7, 10, 11) are generally smooth, although on the protoconch of C. fusiformis, there is a small spiral keel in the middle on the two abapical whorls; similar spiral keels can be found in the protoconchs of Cerithiopsis beneitoi Rolan, Espinosa & Fernandez-Garcés, 2007 and Cerithiopsis dilata Rolan, Espinosa & Fernandez-Garcés, 2007. Olsson & Harbison (1953) described Cerithiopsis aralia, from the Phocene north of St. Petersburg, Florida, and stated that this species should be carefully compared with Cerithiopsis fusiformis. Jong & Coo- mans (1988) considered Cerithiopsis brassica Olsson & Harbison, 1953 as a synonym. We examined the holotype of C. aralia, and although both species are very similar, we prefer to retain them as separate taxa because we feel that additional studies are necessary to confirm the possible synonymy of species from such distant geological provenance. Although we did not examine the holotype of C. brassica, the same interpretation applies. The previously known distribution of C. fusiformis included the Western Atlantic, from North Carolina (U.S.A.) to the Caribbean (Rosenberg, 2005); C. aimen and C. prieguei were known only from Cuba (Rolan & Espinosa, 1995). This is, therefore, their first record from the South Atlantic. In Brazil, C. fusiformis 1s widely distributed, from the states of Pernambuco (northeastern Brazil, ~ 8°S) to Rio Grande do Sul (southern Brazil, = 30°S); C. prieguei and C. aimen have only been found in southeastern Brazil (Espirito Santo State, = 19°S). Both Cerithiopsis balaustium and Cerithiopsis capixaba are restricted to localities in southeastern Brazil: about 24°S and 20°S—24°S, respec- tively, with some records of Cerithiopsis balaustium from the northeast coast, though in a southern locality (=13°S). Acknowledgments. Dr. A. Baldinger (MCZ) for loan of types; Dr. P. Bouchet and Mr. P. Maestrati (MNHN) for loan of several lots from northeast Brazil; Dr. L. Simone (MZSP) for taking the photos of the types; Dr. O. Soriano (MNCN), for loan of types; Dra. C. Miyaji (IOUSP), for loan of material; Dr. J. Harasewich (USNM) for allowing the examination of types; Dr. C. Redfern for comments on the identification of the species and loan of specimens from Bahamas Islands and sending of bibliography; Dr. P. Bouchet (MNHN) for comments on the taxonomy of Cerithiopsis; Dr. E. Rolan (Museo de Historia Natural, Spain), for revision of the manuscript and for sending bibliography; and Dr. R. Absalao (UFRJ), for reading and commenting on the manuscript; Dra. Janet Reid, for revising the english text; Mr. R. Martins (Petrobras Research Center) for taking SEM photos of some shells. REFERENCES Boyko, C. B. & J. R. CORDEIRO. 2001. Catalog of Recent Type Specimens in the Division of Invertebrate Zoology, American Museum of Natural History. Bulletin of the American Museum of Natural History. 262, vol. Mollus- ca, part 2, 170. CLENCH, W. J. & R. D. R. TURNER. 1950. The Western Atlantic Marine Mollusks Described by C. B. Adams. Occasional Papers on Mollusks 1(15):233-403, pls. 28-49. JAY, M. & J. DRIVAS. 2002. The Cerithiopsidae (Gastropoda) of Reunion Islands (Indian Ocean). Novapex 3(1):1-45. JONG, K. M. DE & H. E. COOMANS. 1988. Marine gastropods from Curacao, Aruba and Bonaire Studies on the Fauna of Curacao and other Caribbean Islands. Leiden 69:1— 261, 47 pls. LASERON, C. F. 1951. Review of the New South Wales Cerithiopsidae. Australian Zoologist 11:351—368. LASERON, C. F. 1956. The Family Cerithiopsidae (Mollusca) from the Solanderian and Dampierian Zoogeographical Provinces. Australian Journal of Marine and Freshwater Research 7:151—182. MARSHALL, B. A. 1978. Cerithiopsidae of New Zealand and a Provisional Classification of the Family. New Zealand Journal of Zoology 5:47—120. OpvgE, H. 1989. Distribution and records of the marine Mollusca in the northeast Gulf of Mexico—a continuing monograph. Texas Conchologist 26(1):10—30. Ousson, A. A. & A. HARBISON. 1953. Pliocene Mollusca of Southern Florida, with special reference to those from North Saint Petersburg. Monographs of the Academy of Natural Sciences of Philadelphia 8:vii + 459 p., 65 pls. PONDER, W. F. 1998. Prosobranch classification. In: P. L. Beesley, G. J. B. Ross & A. Wells (eds.), Mollusca: The Southern Syntesis. Fauna of Australia. Vol. 5, Pp. 566— 568, CSIRO Publishing, part B. viii, 565—1234. PONDER, W. F. & A. WAREN. 1988. Classification of the Caenogastropoda and Heterostropha - a list of the family- group names and higher taxa. 288—328, In: W. F. Ponder (ed.), Prosobranch Phylogeny — Proceedings of a symp- sium held at the 9th International Malacological Con- gress, Edinmbug, Scotland. Malacological Review Sup- plement 4. Ann Harbor. 346 pp. REDFERN, C. 2001. Bahamian Seashells. A Thousand Species from Abaco, Bahamas. Bahamianseashells.com_ Inc.: Boca Raton. 221 p., 124 pls. ROLAN, E. & J. ESPINOSA. 1995. The family Cerithiopsidae (Mollusca: Gastropoda) in Cuba 3 The genus Cerithiopsis s.l., species with brown shells. Iberus 13:129-147. ROLAN, E., J. ESPINOSA & FERNANDEZ-GARCES. 2007. The Page 80 family Cerithiopsidae (Mollusca: Gastropoda) in Cuba 4. The genus Cerithiopsis s.1., the banded and the variably coloured species. Neptunea 6(2):1—129. ROSENBERG, G. 2005. Malacolog 4.0: A database of Western Atlantic marine Mollusca. [WWW database (version 4.1.0)] URL http://data.acnatsci.org/wasp]. The Veliger; Vol. 50; NorZ USTICKE, G. & W. NOWELL. 1959. A Check List of Marine Shells of St. Croix., vi + 90p., 4 pls. VOKES, H. E. & E. H. VOKES. 1983. Distribution of Shallow- Water Marine Mollusca, Yucatan Peninsula, Mexico. Middle American Research Institute: New Orleans. 182 p., 50 pls., 9 figs., 1 map. The Veliger 50(2):81—82 (June 20, 2008) THE VELIGER © CMS, Inc., 2007 A Record of the Invasive Slug Veronicella cubensis (Pfeiffer, 1840) | in California R. J. MC DONNELL,'” A. HANSEN,' T. D. PAINE! AND M. J. GORMALLY° "Department of Entomology, University of California, Riverside, California 92521, U.S.A. (e-mails: rory.mcdonnell@ucr.edu; allison.hansen@email.ucr.edu; tpaine@ucr.edu) ?Applied Ecology Unit, Centre for Environmental Science, National University of Ireland, Galway, Ireland (e-mails: mike.gormally@nuigalway.ie) Abstract. Veronicella cubensis (Pfeiffer, 1840) is reported from California for the first time and the significance of the find in terms of agricultural production and the effectiveness of invasive gastropod screening at air and seaports within the state are discussed. Following a survey of the invasive slug fauna of California, we present the first record of Veronicella cubensis (Pfeiffer, 1840) (Stylommatophora: Veronicel- lidae) on the west coast of North America. A single individual was collected under a potted plant in a garden center in Santa Barbara, California (N34°24”908", W119°45.018”) on 19th June 2006. Other slug species collected in the same location were Deroceras reticulatum (Miller, 1774), Deroceras panor- mitanum (Lessona and Pollonera, 1882), Lehmannia valentiana (d’Audebard de Férussac, 1823) and Arion hortensis d’ Audebard de Férussac, 1819. The specimen has been deposited at Department of Malacology, Academy of Natural Sciences in Philadelphia under catalogue number ANSP-A21201. Although having a highly variable coloration (in- cluding an albino form), the notum of V. cubensis is usually dark to pale brown (Figure 1), generally with a pale dorsomedian line. It often has black speckling which sometimes fuses to form two lateral bands (Robinson and Hollingsworth, 2004). The female genital pore ‘is located closer to the pedal sulcus than the peritoneum. The penis has a characteristic flaring that produces a blade-like structure down each side and the penial gland has numerous, very long tubules (David G. Robinson, pers. comm.). It is thought that V. cubensis is native to Cuba (Robinson and Hollingsworth, 2004) but it has also been introduced to Jamaica (Baker, 1925), Bahamas, Haiti, Dominican Republic, Puerto Rico, Guam (Thomé, 1993a), Antigua, Saint Kitts and Nevis, Dominica, Barbados (West Indies), St. Coix, Olosega (Manu ‘u Islands), Tutuila (American Samoa), Pohnpei (Micro- nesia), Rota, the Northern Mariana Islands (Robinson and Hollingsworth, 2004) and Hawai'i (Thome, 1993b). It has also been previously collected (interceptions) in Florida and New Orleans (Thome, 1993a). In 2002, V. cubensis was listed as the seventh most potentially damaging gastropod of either agriculture or natural ecosystems if it became established in the U.S. (Cowie, 2002). Since this species was introduced into Hawai 1in 1985, it has caused severe damage to vegetable, ornamental, and landscape plants, and the species is now a potential threat to Hawar’i’s $104 million vegetable and floriculture industry (Hata et al., 1997). The species 1s also an extremely serious agricultural and horticultural pest on Rota and Guam (Robinson and Hollingsworth, 2004). In addition, V. cubensis has been associated with the transmission of the trematode Angiostrongylus cantonensis (Chen) which causes the potentially lethal eosinophilic meningitis in humans (Cuba: Aguiar et al., 1981; Jamaica: Lindo et al., 2004) Although our record is of a single specimen and return trips to the same location and surrounding areas on 2 August 2006 and 14 March 2007 did not yield any additional individuals, our discovery 1s still of concern as it indicates that some potentially severe pest gastropods are not being detected during pre-clearance at US. seaports, airports and border crossings. Such failed interceptions represent the first step in establishment of exotic species in the U.S. and as such these finds should be reported as they may help to prevent further invasions and the ultimate establishment of pestiferous species. Although some species may not be of concern at their port of entry, they represent a source for transport to other areas of the U.S. and the world, where they could potentially become established and become serious pests. In the case of V. cubensis, spread of individuals to areas such as New Orleans could result in such a scenario. In addition, V. cubensis is predomi- nantly a tropical to subtropical species (Gomes and Thome, in press) and heavily-irrigated desert areas in southern California, including extensive and diverse urban landscape environments, may be a vulnerable habitat for colonization, as they are likely to provide the hot, humid conditions favored by tropical gastropods Page 82 oo ~ The Veliger; Vol: 50, Now Figure 1. Specimen of Veronicella cubensis (Pfeiffer, 1840) collected in Santa Barbara, California. such as V. cubensis. It is therefore imperative that improved screening of shipments that are known to harbor invasive slugs and snails e.g., tiles, fruit, vegetables and ornamentals (Robinson, 1999) are put in place to help mitigate the problem of invasive gastropods in the U.S. Acknowledgments. Sincerest thanks to David Robinson (USDA) for confirming the identification ,of the specimen and for permission to use unpublished data. Thanks to Gary Bernon (USDA) for an English translation of Maceira (in press). We are grateful to G. Bernon, J. Harwood, R. Hollingsworth, D. Robinson and C.D. Williams for review comments. This research was funded by the European Union under a Marie Curie Outgoing International Fellowship (MOIF-CT-2005-21592). LITERATURE CITED AGUIAR, P. H., P. MORERA & J. PASCUAL. 1981. First record of Angiostrongylus cantonensis in Cuba. American Journal of Tropical Medicine & Hygiene 30(5):963—965. BAKER, H. B. 1925. North American Veronicellidae. Proceedings of the National Academy of Sciences, Philadelphia 77:157-84. Cowlk, R. H. 2002. List of potential pest mollusks in the U.S.A.. Interim Report to USDA-APHIS-PPQ. 8 pp. Gomes, S. R. & J. W. THOME. In press. Diversity and distribution of the Veronicellidae (Gastropoda: Soleoli- fera) in the Oriental and Australian biogeographical regions. Memoirs of the Queensland Museum. HATA, T. Y., A. H. HARA & B. K.-S. Hu. 1997. Molluscicides and mechanical barriers against slugs, Vaginula plebeia Fischer and Veronicella cubensis (Pfeiffer), (Stylommato- phora: Veronicellidae) Crop Protection 16(6):501—506. LINDO, J. F., C. T. ESCOFFERY, B. REID, C. CODRINGTON, C. CUNNINGHAM-MyRIE & M. L. EBERHARD. 2004. Fatal autochthonous eosinophilic meningitis in a Jamaican child caused by Angiostrongylus cantonensis. American Journal of Tropical Medicine and Hygiene 70(4):425-428. ROBINSON, D. G. 1999. Alien invasions: the effects of the global economy on non-marine gastropod introductions into the United States. Malacologia 41(2):413-438. ROBINSON, D. G. & R. G. HOLLINGSWORTH. 2004. Survey of slug and snail pests on subsistence and garden crops in the islands of the American Pacific: Guam and the Northern Mariana Islands. Part 1. The leatherleaf slugs (Family Veronicellidae). Internal Report for U.S. Department of Agriculture and the Government of Mariana Islands, 11 pp. THOME, J. W. 1993a. Erneute Beschreibung von Veronicella cubensis (Pfeiffer 1840) (Gastropoda:). Archiv fuer Molluskenkunde 122:113—121. THOME, J. W. 1993b. Estado actual da sistematica dos Veronicellidae (Mollusca; Gastropoda) americanos, com comentarios sobre sua importancia economica, ambiental e na satide. Biociencias, Porto Alegre 1(1):61—75. Personnal Communications: (1) David G. Robinson, USDA APHIS PPQ, Depart- ment of Malacology, Academy of Natural Sciences, 1900 Ben Franklin Parkway, Philadelphia, Pennsylva- nia 19103, U.S.A. THE VELIGER ) yy) The Veliger 50(2):83-96 (June 20, 2008) © CMS, Inc., 2007 Developmental Mode in Opisthobranch Molluscs from the Tropical Eastern Pacific Ocean JEFFREY H. R. GODDARD Marine Science Institute, University of California, Santa Barbara, CA 93106-6150 (e-mail: Goddard@lifesci.ucsb.edu) ALICIA HERMOSILLO University of Guadalajara, Centro Universitario de Ciencias Biol6gicas y Agropecuarias, Km 15.5 Carretera a Nogales, Zapopan, Jalisco, México Abstract. Little has been published on mode of development in benthic opisthobranchs from the tropical eastern Pacific Ocean. Based on observations of uncleaved eggs, developing embryos, or hatching larvae, we determined or inferred mode of development for 43 species collected primarily from Bahia de Banderas, México. Forty-two of these, including the umbraculoidean Ty/odina fungina, had planktotrophic development, while Phidiana lascrucensis hatched as lecithotrophic larvae. Both the sacoglossan Elysia pusilla, which had small eggs, relatively large egg capsules, and irregular strands of extra-capsular yolk in its egg mass, and the dendronotacean Lomanotus vermiformis may also have lecithotrophic development in this region. Combined with previously existing data, mode of development is now known for 91 species of native, benthic, shallow-water opisthobranchs from the eastern Pacific and can be tentatively inferred for another 13 species based on data from other regions. Four species hatch as lecithotrophic larvae, and the remaining 100 as planktotrophic larvae. The prevalence of planktotrophic development in opisthobranchs from the eastern Pacific is similar to that known from the adjacent northeast Pacific Ocean, but is higher than in the less productive waters of the Caribbean Sea and the tropical western Atlantic Ocean, where opisthobranch eggs attain much larger diameters and 37% of the 112 species examined have either lecithotrophic or direct development. These results agree with those known for a diverse range of marine invertebrates across the Isthmus of Panama and are consistent with evolutionary trends expected in the egg size and mode of development with historical changes in ocean productivity and larval feeding environment. INTRODUCTION greater tropical western Atlantic Ocean (together referred to hereafter as the W Atlantic). Based on the distributions reported recently by Behrens & Hermosillo (2005), Camacho et al. (2005) and Hermosillo et al. (2006), 17 species included in Goddard (2004) have ranges extending well into the Panamic province. Excepting Antaeolidiella indica (Bergh 1888), which has lecithotrophic development and is circumtropical in distribution, all have plankto- More than 90% of 126 native species of opisthobranchs studied from the cool temperate waters of the northeast Pacific Ocean have planktotrophic development, a high prevalence thought to reflect the region’s suitability for larval feeding and growth, especially its generally slow currents, large geographic expanse, and primary production driven by coastal upwelling and horizontal advection via the California Current, (Goddard, 2004). In contrast, relatively little information has been published on developmental mode in opisthobranchs from the tropical eastern Pacific (hereafter E Pacific), which includes the Panamic biogeographic province and extends from southern Baja California Sur to central Peru (Briggs, 1974). Here we document developmental mode in 43 species of shallow-water opisthobranchs known from the Pacific coast of Mexico, and compare the frequencies of the major modes of development to those known from the NE Pacific as well as the neighboring Caribbean Sea and trophic larval development. Gonsalves-Jackson (2001, 2004) compared development in opisthobranchs across the Isthmus of Panama, and observed “planktonic” development in all 39 Pacific species she studied. Although Gonsalves-Jackson (2004) did not distinguish between planktotrophy and pelagic lecithotrophy, the small egg-sizes she reported (under 70 microns for sacoglossans and under 115 microns for all other species), combined with her descriptions and illustra- tions of embryos and hatching larvae, reliably indicate planktotrophic development for all 39 species, based on morphological criteria and known egg-size distribu- Page 84 The Veliger, Vol; 50) Now! Nayarit Punta Mita Islace ein Marietas Bahia de Banderas {indomar Los Arcos Mismaloya ——— Jalisco x Pacific Ocean Figure 1. Map of Bahia de Banderas, México, showing collection localities. tions (Thompson, 1976; Bonar, 1978; Hadfield & Miller, 1987; Goddard, 2004). Finally, information on the development of 13 of the more widely distributed Panamic opisthobranchs has been obtained from other regions of the world. Owing to the possibilities of geographic divergence and poecilogony, these latter data cannot be considered definitive for the E Pacific, but do suggest that Lomanotus vermiformis Ehot 1908 and Phestilla lugubris (Bergh 1870) have lecithotrophic development, and that the remaining 11 species are planktotrophic (Harris, 1975; Bandel, 1976; Clark & Goetzfried, 1978; Switzer-Dunlap, 1978; Schmekel & Portmann, 1982; Gonsalves-Jackson, 2004: Table 1). Taken together, these data suggest that the prevalence of planktotrophy in the E Pacific is similarly high to that observed in the NE Pacific. To better compare developmental mode in opistho- branchs from the two regions, we present here data on the development of 45 species of opisthobranchs from the Panamic biogeographic province. We then compare our results from the E Pacific to those from the more seasonally stable and relatively oligotrophic waters of W Atlantic, where a shift toward larger egg sizes and benthic or non-planktotrophic modes of development has been documented in opisthobranch gastropods (Gonsalves-Jackson, 2001, 2004), as well as other taxa, including bivalve molluscs, crustaceans, echinoid echi- noderms, and bryozoans (Lessios, 1990; Jackson & Herrera, 1999; Marko & Moran, 2002; Wehrtmann & Albornoz, 2002: Fortunato, 2004; Moran, 2004). During our survey, we were able observe the egg masses and hatching larvae of the umbraculoidean opisthobranch Tylodina fungina Gabb, 1865. Because so little is known about the development and larvae of umbraculoideans (see Gibson, 2003), we also provide here a detailed description of its egg mass and hatching larvae. METHODS Adult opisthobranchs and opisthobranch egg masses positively identified in the field were collected by hand from 16-28 February 2006 from subtidal and intertidal sites within Bahia de Banderas (20°30’N, 105°30’W), in the states of Jalisco and Nayarit, on the west coast of Mexico (Figure 1). This bay, approximately the size, shape and depth of Monterey Bay, California, and the collecting localities, have been described by Hermosillo (2003). Local sea surface temperatures during our work period averaged approximately 21°C. Adults were held in the field laboratory in containers (250 to 2000 ml) of seawater at 19—-23°C until they laid egg masses. Recently laid egg masses were examined using a compound microscope equipped with an ocular micro- meter. If first cleavage had not commenced, the diameters of a random sample of 10 zygotes were measured in each egg mass; otherwise, an upper limit on zygote size was estimated by measuring the dimensions of a few randomly selected embryos at or before the gastrula stage and (or) the minimum dimension of tightly fitting egg capsules containing embryos at or before the gastrula stage of development. After initial examination, each egg mass (or a portion of a large egg mass) was then isolated in a separate, labeled vial. The seawater in these vials was changed J. H. R. Goddard & A. Hermosillo, 2007 daily, and the egg masses examined daily for hatching. Hatching larvae were then examined, measured and in some cases photographed. Developmental mode (planktotrophic, lecithotrophic and direct) and larval shell type (coiled type 1 and egg-shaped, inflated type 2) were assigned according to the egg size distributions and larval morphological criteria described by Thomp- son (1961, 1976), Bonar (1978), Clark & Jensen (1981), Hadfield & Switzer-Dunlap (1984), Hadfield & Miller (1987) and Goddard (2004). After obtaining the above egg masses, most of the adult specimens were relaxed in 7.5% MegCh, fixed in 70% ethanol, and deposited as voucher specimens in the California Academy of Sciences. Other adults were returned alive to the field. We used an underwater data logger (StowAway Tidbit, Onset Computer Corp.) to record temperature at 10 min. intervals in our holding containers. We estimated egg size for Polycera alabe (COLLIER & FARMER 1964) and late embryo size for Lomanotus vermiformis, by placing a mm scale bar next to egg masses in the field and taking underwater digital images. Additional data, obtained by the senior author in central and southern California and Baja California, (1) supplement the data for three species we collected in Bahia de Banderas, and (2) are given for six additional species whose geographic ranges include the Panamic province. To compare frequencies of development modes in the E Pacific to those in the NE Pacific and the W Atlantic, we assigned developmental mode (according to the criteria mentioned above) using published data on species from the E Pacific, and then combined these results with our own. We then calculated the frequen- cies of the different modes of development for (1) the NE Pacific based on Goddard (2004, 2005), and Krug et al. (2007); and (2) the W Atlantic based on data and determinations of developmental mode in Bandel (1976), Clark & Goetzfried (1978), Eyster (1980, 1981), Clark & Jensen (1981), DeFreese & Clark (1983), Carroll & Kempf (1990), Cronin et al. (1995), Ortea (2001), Gonsalves-Jackson (2004), and Pierce et al. (2006). For 17 species from the W Atlantic we assigned mode of development as either planktotrophic or direct based on close-up images of egg masses with either small or large eggs/embryos in Valdés et al. (2006); our determinations for six of these species were confirmed by information in the other references cited above for the W Atlantic, giving us confidence that our determinations for the other 11 species (for which no other information exists) were accurate. Ortea (2001) listed egg sizes for 11 Caribbean species of the nudibranch genus Doto, but provided no information on hatching stages or type of development. For six of these species we assigned developmental mode based on the egg-size distributions known for each mode, conservatively considering species with eggs less than Page 85 100 um in diameter as planktotrophic, species with eggs greater than 165 um diameter as lecithotrophic, and those with eggs greater than 220 um as direct developers. To avoid tabulating Atlantic species more than once, we used Valdés et al. (2006) and the Sea Slug Forum (http://www.seaslugforum.net/) to check for synonymies and recent taxonomic revisions. We used JMP (version 7.0, SAS Institute, Inc.) to (1) conduct contingency analyses of the frequencies of planktotrophic vs. non-planktotrophic (lecithotrophic and direct) development by region, and (2) compare egg size distributions by region. For the latter we excluded egg size data obtained outside the regions of interest, but used all other determinations of egg size available from the above sources, even if mode of development is unknown. Owing to (1) the large number of species of opisthobranchs from the W Atlantic found to have non-planktotrophic develop- ment, and (2) the lack of detailed phylogenies for most opisthobranch taxa across the Isthmus of Panama, we limited the contingency analysis of developmental mode by E Pacific vs. W Atlantic Oceans to the numbers of families, rather than species, with plankto- trophic vs. non-planktotrophic development. To reduce the influence of phylogenetic constraints unique to particular families, we further limited this analysis to families common to both oceans. RESULTS We found 70 species of opisthobranchs in Bahia de Banderas and obtained data on the development of 39 of these. Combined with the data on six Panamic species obtained by the senior author outside of Bahia de Banderas, data on development were obtained for a total of 45 species (Table 1). Forty-two had plankto- trophic development, and the aeolid nudibranch Phidiana lascrucensis Bertsch & Ferreira 1974 had lecithotrophic development (Table 1). Although we obtained data on the egg and embryo size, respectively, of the sacoglossan Elysia pusilla (Bergh 1872) and dendronotacean nudibranch Lomanotus vermiformis (Table 1), we did not observe their hatching stages and were unable to determine with certainty their mode of development. The former had small eggs but had relatively voluminous egg capsules surrounded by irregular strands of extra-capsular yolk (ECY) (Fig- ure 2). As shown by Clark & Jensen (1981), the presence of these traits can indicate lecithotrophic or even direct development in sacoglossans, despite otherwise small egg size. The embryos of Lomanotus vermiformis observed in the field were large enough (Table 1) that none of the three major modes of development can be ruled out based solely on known distributions of egg size and shell size at hatching (Hadfield and Miller, 1987; Goddard, 2004). 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Ay ore yoy “(epnsyses Yysno1yy [[99 91) soJOSAZ YIM UT Sased UT “sosoyjuored ur ozis ojdures ‘S.17 Ajoveurxoidde 0 1¥ soinjesoduia} aovyj.ns-ves ‘potiod MUOAIQUID SY} JO [[B IO SOUL 1OF PfOY SosseUu 389 10} ATUO UDdAIS mesodway (SpoysA 99S ‘SUOT}BATOSGO jeuosiad) sojOsAzZ 9Y} UY} WOTSUSUIpP ysoyeoIs UT JOSUOT IIHT P so8vys SUOKIQUUS AJR JO S}USUTDINSPIUT UO Poseq ‘UDAIS SI OZIS S39 1OJ WUT] raddn uv ‘paasosqo JOU 310M YUM “UONVIASp plepuRys 9UO SNUTU 10 snjd ‘suevow oie Surysyey 1e y {Busy [Joys pue 1oyourvrp 339 JO} sanyeA ‘sosoyyuared ur oie juowidoyaaap Jo sopour patsajuy “(PO0T “PABPPOD 398) SIQUISUOD YIM SUOSLIeduIOS pue ‘yuourdoyaaop Jo sopour 1ofew sy} 1OJ (L861) TNA PUP PISYPPH Jp (orydosjoy use] “7 corydonoyyqueyd “gq su ‘QsIMIOY}O payloads ssapu “Psye[Oo Pep CU sSuruasoidot kq paviodas suonnqtiystp ozis 389 94} UO peseq Pottojut Sem JUdWIdOJSAIP JO POUL “PAAtIsgO JOU IJOA\ OVATE] Suryovey audopaaap Jo 9poy “(] P4NST 99S) COINS ‘WIeAVN pure oosies ‘sesopurg op eye Ul oe SoTpLIOT [Ie (—) ysep & YUM ‘Aroye1OQR] 94} UT pousodap 10 poly oY} UF poyda][O9 Joy “ssvul S30 g[SuUIS B WO. poure}qo sem vIeP JO MOI YOR” “ULIDO IYOR_ W19ISvO jeoido. oy} WO1y UMOUY syourrqoyjsido ur yuotudoarop SIUOAIQUID TO PIEP sareredwo) 1 19 Page 87 J. H. R. Goddard & A. Hermosillo, 2007 IVUlOpUrT] sPIULOFIeD eleg ‘olsesoy vjung [ePHAaUI ‘sooIW SOT vAOTRUISTI Ievwopury vAOTRUISTI selopurg op viyeg VIUIOJILO ‘ye 9181S O1M op vurUO/y [eplso}UrI ‘sooly soy Ieulopury PAOTRUISTIA [BPHAa UI ‘sory SOT SPOIL SPIOLIVIA| svlopurg op viyeg PAOTRUISTIA] BIUIOFILD vleg ‘oVliesoy ejung Ieulopury PU eUNg PAOTRUISII rewopury seUIOM[LS eleg ‘oVIesoy viung Aylpeso07 oT) d d yUOUU -dojaaop JO Spo ou ou ou ou ou sulyo}ey ye sjodsaAq (Ol) Tr + 8 9LT 061 ~ (07) O€ = 6THT (7) 81 + €'rOl (9) 88 + PLTI (Ol) 9b = VITI SBll= (Ol) (t+ VLTI ()QLps (OI) ST + VP 67TI (Ol) 61 + S*€0I (OI) L's + 9'6EI (um) Suryo}ey ie yisug] []oys I 6-9 s9 I ed — I tc-6l iS c LI-Cl Ol I oie = if — —— I tc-6l Si I €c-6l =< I €c-6l 9 I col S I tc 6l L I tc 6l v I 6I-91 SI od4} = (2D) (sAep) wus ‘dwoy porsod o1uo0AIQuig ‘ponunuodg T 91981. 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(sAep) ‘dup = poriod s1UOAIQUUI ‘ponuluod 1 919PL I I I I ansdeo Jod s330 7 LL> (O01) 81 + 9°94 OSl> blL> (pF) 90 + 199 OOl> (Ol) rl + 9°88 (wm) JoouRIp 33q (€T8I oferyD aJoq) vunjiyodnau nypands TOIPLI “9SOPLI ZISVO rLol BILDIID-] W YOSWOG SIsuaINAISY] DUDIPIYd 880PZI ZISVO €00T vsouldsy wW AI[eQRD ‘vIIO VIDIPLUDUNYDY VIISOUWAIL] 6SOELI ZISVO SOOT O][ISOWIOF] 2 susiysg Jo | ‘ds puyjaqvpy 6SObL1 ZISVO LOG] SNOIVIAL WZ snoiefy V/ja7 vuIyjagvpT 0661 URLIIZN YW AMUSO wWNsLOSNIADLU DUIPJAGP}] O90PLI ZISVO LOOT OSOSUOLL W VIOIVD AVINA[YUAJA SNULIOAD.T PoOrLI ZISVO ‘ds snyoupaqny LSOPLI ZISWO SOOT [Re JO BIDIeH-oyseuvD Jo ZT ‘ds snyouvaqny c90ELI ZISVO SR6] SUDIYIG sMpjNon. snyoUuvAGNT 8SOPLI “8S0EL1 ZISVO €OOT SIPIPA 2P OINSUW anz1] DUoYIND UOX? | J. H. R. Goddard & A. Hermosillo, 2007 Page 89 Figure 2. Elysia pusilla. A. Detail of egg mass, showing egg capsules (C), extra-capsular yolk (ECY), and embryos (E). B. Close-up of extra-capsular yolk on outside of egg capsules. Goetzfried (1978) reported that this species has lecithotrophic development in Florida. We obtained data on the development of nine undescribed species, eight of which are recognized and illustrated in Behrens & MHermosillo (2005), Camacho et al. (2005) or Hermosillo et al. (2006), and one of which (ELubranchus sp.) appears to be new. Our record of Doris immonda (Risbec 1928) from under coral rubble at Punta Mita, the northern boundary of Bahia de Banderas, is the first sighting of this species in Mexican waters and only the second record of its occurrence outside the Indo-west Pacific (Camacho et al., 2005). Voucher specimens for all ten of these species have been deposited in the California Academy of Sciences (Table 1). Combined with previously existing data from the region (see Introduction), mode of development is now known for approximately 91 species of native, benthic, shallow-water opisthobranchs from the E Pacific and can be tentatively inferred for an additional 13 species based on published data from other regions. The aeolid nudibranchs Antaeolidiella indica and Phidiana lascru- censis, and likely also both the dendronotid Lomanotus vermiformis and the aeolid Phestilla lugubris, hatch as lecithotrophic larvae, and the remaining 100 as planktotrophic larvae (Table 2). Fifteen of the 39 E Pacific species studied by Gonsalves-Jackson (2004) were only identified to genus (with voucher specimens of each deposited in the American Museum of Natural History). Depending on the overlap with species we studied, this might reduce the total number of E Pacific species whose development is known, but it wouldn’t significantly affect the overall prevalence (approxi- mately 4%) of lecithotrophic development known from this region. By number of species, the incidence of planktotrophic vs. non-planktotrophic development in the E Pacific did not differ significantly from that observed in the NE Pacific Ocean (Table 2, Likelihood ratio y¥° = 1.735, P = 0.188). From the literature we were able to determine or infer mode of development for 112 species of opistho- branchs from the W Atlantic (Tables 2 & 3). Lecitho- trophic and direct modes of development were more prevalent in the W Atlantic compared to the E Pacific, comprising 21% and 16%, respectively, of the 112 species. In the W Atlantic these non-planktotrophic or non-feeding modes of development occurred in 17 families (Table 3) from all of the major orders and suborders of benthic opisthobranchs, save the Umbra- culida, the development of which has not been examined in the W Atlantic. Sixteen of these families have representatives in the E Pacific, where non-feeding development is known from four and planktotrophic development from all 16 (Table 3). Limiting the contingency analysis to these 16 families, the incidence of non-feeding development, by number of families, is significantly higher in the W Atlantic than in the E Pacific (Likelihood ratio y* = 5.830, P = 0.016). The higher prevalence of non-planktotrophic modes of development in the W Atlantic Ocean, compared to Page 90 The Veliger, Vol. 50, No. 2 Table 2 Number of species of opisthobranch molluscs known or inferred to have planktotrophic, lecithotrophic or direct development in the tropical eastern Pacific Ocean, compared to the NE Pacific and tropical western Atlantic Oceans. Mode of development Region Planktotrophic Lecithotrophic Direct Sources of data NE Pacific 116 4 6 Goddard (2004, 2005) Krug et al. (2007) E tropical Pacific 100 4 0 Harris (1975), Bandel (1976), Clark & Goetzfried (1978), Switzer-Dunlap (1978), Schmekel & Portmann (1982), Gonsalves-Jackson (2004), Goddard (2004), present study W tropical 7) 24 17 Bandel (1976), Clark & Goetzfried (1978), Eyster (1980, 1981), Clark & Atlantic Jensen (1981), DeFreese & Clark (1983), Carroll & Kempf (1990), Ortea (2001), Gonsalves-Jackson (2004), Pierce et al. (2006), Valdés et al. 2006 Table 3 Number of species of opisthobranchs from the tropical eastern Pacific and tropical western Atlantic Oceans known or inferred to have planktotrophic or non-planktotrophic (= lecithotrophic and direct) development, by taxonomic family. Based on sources listed in Table 2; taxonomic classification according to Behrens & Hermosillo (2005) and Valdés et al. (2006). A blank space means that no representatives of that family are known from that ocean. Tropical eastern Pacific Tropical western Atlantic Family Planktotrophic Non-planktotrophic Planktotrophic Non-planktotrophic Aegiretidae 1 0 Aeolidiidae 6 1 Aglajidae 2 0 Aplustridae Aplysiidae 10 Arminiidae 1 Boselliidae Bullidae Caliphyllidae Chromodorididae Conualevidae Corambidae Cylichnidae Dendrodorididae Dorididae 1 Dotoidae Eubranchidae Facelinidae Flabellinidae Goniodorididae Haminoeidae Hancockiidae Hermaeidae Hexabranchidae Limapontiidae Lomanotidae Oxynoidae Placobranchidae Pleurobranchidae Polyceridae Tergipedidae Tritoniidae Tylodinidae Zephyrinidae Www oe So (o) So CON Re RK NN ArReoorrocownv SCNNNANWUNAWNK KOKO oooocoroqocooqocooqocoqcjo OOOrONAYNF RNR OOF COWOA HHO O Ne WN PP WR RR R oocorocoocoocooroe SDGDOrFONAWNDHRONODOWWHWKEWORDHON OF — — & — Total no. of species 100 J. H. R. Goddard & A. Hermosillo, 2007 4+ - 25 15 Ee 5 50 100 150 200 250 300 350 No. of species No. of species 50 100 150 200 250 300 350 Egg diameter (microns) Figure 3. Egg size distributions, with box plots, for Opisthobranchia from the E Pacific (n = 81 species) (A) and the W Atlantic (n = 92 species) (B). The vertical line in the box plots indicates the sample median, and the diamond indicates the sample mean and 95% confidence intervals. The egg size distribution for the E Pacific is based in part on the upper size limits presented for some species in Table 1. The actual egg sizes for these species are slightly smaller (see heading for Table 1). the E Pacific, is reflected in the strong skew toward larger egg sizes in the former (Figure 3). DISCUSSION Planktotrophy is the dominant mode of development in opisthobranchs from the E Pacific, as in the NE Pacific. Differences in sea surface temperatures aside, both regions have productive waters seemingly conducive for larval feeding and growth. Seasonal, wind-induced, coastal upwelling and the slow-moving, nutrient-rich, California current fuel primary production in the NE Pacific (Bernal & McGowan, 1981; Chelton et al., 1982; Bakun, 1990; Mann & Lazier, 1991), while three types of upwelling (wind-induced coastal, equatorial, and that associated with the cyclonic gyre known as the Costa Rica dome) fuels production in the E Pacific (Wyrtki, 1964; McCreary et al., 1989; Mann & Lazier, 1991; Fielder, 1992). Larval food supplies in both regions are therefore probably rarely limiting, contrib- Page 91 uting to the evolutionary maintenance of small egg sizes and planktotrophy in taxa not _ historically constrained to non-feeding modes of development. The lack of directly developing opisthobranchs in the E Pacific stands in contrast to the NE Pacific, where it has so far been documented in 6 species (Table 2). However, development has been examined in less than less than half of the total number of species of opisthobranchs known from the E Pacific (see Cama- cho et al., 2005; Hermosillo et al., 2006), and sampling in the E Pacific has been biased toward outer coast habitats and, in our study, the winter season. Addi- tional sampling, including in estuarine habitats and during the summer (when a different complement of warmer water species may be present), may therefore be needed to determine if the observed difference in the frequency of this mode of development is significant. Although direct development is not yet known for any opisthobranch from the E Pacific, we suspect that Chromodoris sp. 1 of Hermosillo et al. (2006) may hatch as juveniles, based on its small adult size (10 mm) and a known geographic distribution limited to the Revillagigedo Islands, a small volcanic archipelago located 720 km west of mainland Mexico. Eight of the species whose development we examined in this study were also identified and studied by Gonsalves-Jackson (2004) in Panama. In all cases our size measurements of eggs and embryos corresponded closely with her measurements of egg size, and our observations and measurements of hatching larvae were consistent with her sketches of embryos, only some of which depicted embryos near hatching (Gonsalves-Jackson did not provide measurements of shell size at hatching for any of the species she examined). The largest discrepancy in egg size was for the aeolid Flabellina marcusorum. Although we record- ed its early embryos as being 73 um in largest diameter (indicating a slightly smaller egg diameter), and Gonsalves-Jackson (2004) reported a mean egg diam- eter of 81.5 um, this difference is within normal intra- specific variation, especially between different popula- tions (e.g., Goddard, 1984, 2004; Todd et al., 2001). Planktotrophy was significantly more prevalent in opisthobranchs from the E Pacific than in the W Atlantic, consistent with patterns in egg size and (or) mode of development documented by other workers for prosobranch gastropods, bivalve molluscs, alpheid crustaceans, bryozoans, echinoid echinoderms and reef-forming corals across the Isthmus of Panama (Lessios, 1990; Jackson & Herrera, 1999; Marko & Moran, 2002; Wehrtmann & Albornoz, 2002; Fortu- nato, 2004; Moran, 2004). Most of these studies intentionally compared life history traits in sister species thought to have diverged as a result of the rise of the Isthmus of Panama, thereby ruling out phylogenetic constraints as the sole determinant of the observed Page 92 The Veliger, Vol. 50, No. 2 geographic patterns in developmental mode. Larger egg-sizes and non-feeding modes of development are therefore thought to have evolved in these taxa as a result of environmental factors, namely the drop in productivity of surface waters in the Caribbean Sea and W Atlantic following the rise of the Isthmus of Panama (Bishop & Marra, 1984; Coates & Obando, 1996; Collins, 1996; Allmon, 2001). In this environment, larger eggs (with their greater yolk reserves) might be expected as one mechanism for offsetting the mortality caused directly or indirectly by lower ocean productiv- ity and a poor larval feeding environment (e.g., Vance, 1973; Lessios, 1990). However, evidence presented by Moran (2004) for planktotrophic arcid bivalves, sug- gests that selection for reduced egg sizes in the E Pacific (in response to increased productivity following the rise of the Isthmus) may have been more important in shaping patterns of egg size across the Isthmus in that taxon. Moran (2004) also rightly notes that differential extinction of species with large eggs in the E Pacific and species with small eggs in the W Atlantic following the rise of the Isthmus might also be important in explaining recent egg size patterns. Environmental considerations aside, phylogenetic constraints do not appear to be important in explaining the difference in mode of development observed in opisthobranchs across the Isthmus of Panama. Plank- totrophy is known from nearly all of the 17 families with non-planktotrophic representatives in the W Atlantic and dominates the 16 of those same families that also occur in the E Pacific (Table 3). Given its widespread distribution among even higher taxonomic levels of opisthobranchs in the W Atlantic, non-planktotrophic development appears to have evolved independently in numerous lineages of opisthobranchs in this region. The differences in ocean productivity and other environmental factors mentioned above apply on even broader geographic scales, and appear to be reflected in ocean basin-wide patterns of egg size and developmen- tal mode in shallow water nudibranchs, the most species rich group of opisthobranchs (Goddard 1992, in preparation). In particular, data we have presented here support the hypothesis that eastern ocean regions, with their widespread upwelling, productive waters, and slow boundary currents (e.g., Mann & Lazier, 1991), will tend to maintain a higher frequency of planktotrophic development compared to western ocean regions, which have less productive waters at mid to lower latitudes and faster boundary currents, which might increase the risk of advection of larvae away from favorable settlement sites. Notes on individual species Lomanotus sp. 1 hatched as small, transparent, planktotrophic larvae with a coiled, type 1 shell 117 um long (Table 1). Because shell type is family- specific (Thompson, 1976; Goddard, 2004), the state- ment, without measurements, by Clark & Goetzfried (1976) that L. vermiformis (as L. stauberi) has an inflated, type 2 shell requires confirmation, especially given conflicting reports of shell type in the family. Thompson (1961), relying on Pruvot-Fol’s (1954, fig. 142g) illustration of a coiled larval shell of L. genei Vérany 1846 from Europe, listed this species as having a type | shell. However, Thompson & Brown (1984) characterized the family Lomanotidae as having a type 2 shell, presumably based on Clark & Goetzfried’s (1976) report for L. vermiformis. Tylodina fungina Gabb 1865 The egg ribbons of Tylodina fungina were observed on the surface of its yellow, keratose sponge prey, identified in Bakus & Abbott (1980) as Aplysina fistularis (Pallas 1766) (= Verongia thiona de Lauben- fels 1930). The egg masses were laid flat, often in overlapping, convoluted layers attached primarily to spongin fibers exposed by the grazing activity of adults (Figure 4A & B). The egg ribbons were similar in appearance to those of T. perversa (Gmelin 1791) and T. corticalis Tate 1889 known from E Australia and the Mediterranean Sea, respectively (Thompson, 1970; image in Poddubetskaia, 2002). The ribbons, though thin, were stiffer than those of most other opistho- branchs (personal observations), and appeared to be reinforced by transverse internal walls (Figure 4B). Egg masses laid on the sponge accumulated minute, golden brown, refractile bodies | to 5 um in greatest dimen- sion (Figure 4C & D, compare to 4B). These bodies were likely unicellular cyanobacteria known to be associated with the surface layers of species of Ap/ysina (Maldonado & Young, 1998; Friedrich et al., 1999; Becerro et al., 2003; Usher et al., 2004). They did not occur inside the egg capsules of JT. fungina, and appeared to accumulate in surface folds and furrows (Figure 4C & D). The refractile bodies gave the egg masses an opaque, pale yellow to pale orange-brown appearance and were often dense enough to obscure views of the embryos. In 70% ethanol the egg masses, like the adult slugs, turned deep purple, indicating the presence of uranidine, a pigment sequestered by species of Tylodina from their sponge prey (Teeyapant et al., 1993). Near hatching, the embryos fit very tightly within their egg capsules, and the egg masses did not appear to break down as quickly as observed in more gelatinous opisthobranch egg masses. Hatching larvae lacked both eyespots and propodium, but had a compact, lipid-rich viscera, a distinctive, burgundy-colored mantle organ, and an operculum (Figure 4E). Their type 1 shells averaged 126 um in length (Table 1) and consisted of J. H. R. Goddard & A. Hermosillo, 2007 Page 93 Concentrations of 4 aa ee > Refractile refraeule bodies @ __ bodies Pigmented Shell mantle organ es 7 Velar ea Foot Viscera E Statolith Z. 25 um Operculum Figure 4. Tylodina fungina. A. Egg ribbons laid on spongin fibers and remains of prey sponge, Aplysina fistularis. B. Egg ribbon laid on the dorsal surface of the shell of another Tylodina fungina. Note the relative transparency of this ribbon compared to those laid on A. fistularis. C. Piece of egg mass removed from its sponge substratum, showing concentrations of minute refractile bodies on surface. D. Higher magnification view of minute refractile bodies on surface of egg mass. E. Newly hatched veliger larva, right lateral view. F. Apex of an adult shell (21.3 mm long), showing the protoconch (= embryonic and larval shell), left lateral view, and juvenile shell. Specimen from Bird Rock, La Jolla, California, 12 December 2004. Page 94 The Veliger, Vol. 50, No. 2 about two-thirds of a whorl. As observed in pleuro- branchs, but not most other hatching planktotrophic opisthobranchs (Gibson, 2003), the mantle was not folded over the edge of the shell. We did not observe vigorous swimming by the newly hatched larvae. The protoconch of an adult Tylodina fungina collected by the senior author at La Jolla, California measured 352 um in length and consisted of about 1.5 whorls (Figure 4F), indicating that the larvae of this species grow significantly in the plankton. Erosion of the outer layer of the protoconch was evident, but there did not appear to be any demarcation between the embryonic and larval shell (Figure 4F). Umbraculum umbraculum (Lightfoot 1786) is the only other umbraculoid opisthobranch whose hatching larvae have been described. Like Tylodina fungina, the larvae of this species also hatch with coiled shell, a pigmented mantle organ, and an operculum (Oster- gaard, 1950; Hartley, 1964). The egg masses, embryo size, and protoconch of T. perversa from the Mediter- ranean Sea appear very similar to those of T. fungina (Valdés & Lozouet, 2000; Poddubetskaia, 2002). Tylodina corticalis from eastern Australia lays egg masses similar to those laid by 7. fungina, but with eggs 98 um in diameter (Thompson, 1970) may have lecithotrophic development. The small size of the hatching larvae, their lack of both eyespots and propodium, and the size of the protoconch on the adult shell, all indicate that the larvae of Tylodina fungina are planktotrophic. Howev- er, the viscera developed in a compact arrangement reminiscent of lecithotrophic larvae, and the newly hatched larvae did not appear to be strong swimmers. It would be interesting to know if recently hatched larvae remain in the vicinity of the parental egg masses and consume the microbes associated with the egg masses and underlying sponge. 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Fishery Bulletin U.S. 63:335-372. re The Veliger 50(2):97—106 (June 20, 2008) THE VELIGER © CMS, Inc., 2007 Three New Buccinid Species (Gastropoda: Neogastropoda) from Chilean Deep-Water, Including One from a Methane Seep KOEN FRAUSSEN Leuvensestraat 25, B-3200 Aarschot, Belgium (e-mail: koen.fraussen@skynet.be) JAVIER SELLANES Universidad Catolica del Norte, Facultad de Ciencias del Mar, Larrondo 1281, Coquimbo, Chile (e-mail: sellanes@ucn.cl) Centro de Investigacion Oceanografica en el Pacifico Sur-Oriental (COPAS), Universidad de Concepcion, Casilla 160-C, Concepcion, Chile Abstract. Three deep water species from off the Chilean coast are described as new. Aeneator prognaviter sp. nov. (off Antofagasta) is compared with A. Joisae Rehder, 1971 and A. castillai McLean & Andrade, 1982. The peculiar Aeneator portentosus sp. nov. (off Coquimbo) has the generic placement based on conchological characteristics and is compared with Americominella duartei Klappenbach & Ureta, 1972. Kryptos explorator sp. nov. (off Concepcion) is compared with K. koehleri (Locard, 1896), the generic placement based on conchological characteristics (protoconch and slightly broader peripheral spiral interspace) and radular morphology. Kryptos explorator sp. nov. has been collected at a recently discovered methane seep area off Concepcion (~36°S), but its degree of association to seep fauna is still uncertain. Key Words: Gastropoda, Buccinidae, Aeneator, Kryptos, new taxa, methane-seeps, bio-diversity, East Pacific, Chile. INTRODUCTION The coastal zone off north to south-central Chile, strongly influenced by wind-driven upwelling, is one of the areas with the highest known primary production rates worldwide (Daneri et al., 2000). Consequently this area of the south-eastern Pacific Ocean harbours a vast pelagic and benthic biomass. However, in spite that the benthic fauna has been proven to be rich in endemic species, of which many are still undescribed or unknown, its scarce knowledge still precludes research- ers have an accurate assessment of the diversity along the Chilean margin. The existing literature on benthic communities along the continental margin off north to south-central Chile is restricted mostly to the shelf and upper slope (e.g., Gallardo, 1963; Brattstr6m & Johanssen, 1983). Except for the general results of the R/V Anton Bruun cruise in the Southern Pacific (Garth & Haig, 1971; Menzies et al., 1973), the Russian Expeditions (Mironov & Rudjakov, 1990, and references therein) and general studies of the archibenthal fauna (Andrade, 1986, 1987), there are no detailed studies on bathyal benthic communities. Regarding mollusks, the offshore continental slope and the deep ocean floor were also largely underesti- mated in the past, if not miserably ignored by collectors and malacologists. Today we know that the Chilean coastline and adjacent continental slope harbors many species of molluscs. The result of continuous research conducted by scientific expeditions (from the Lund University Chile Expedition in 1948-1949 to the PUCK-156 expedition in 2001, to mention a few), by local trawlers (McLean & Andrade, 1982) and from shrimpers of the former Soviet Union (Fraussen & Hadorn, 2000; Poupin, 2003), have produced notewor- thy contributions. Most recently, the existence of methane seepage and associated chemosynthetic com- munities in the bathyal zone off central Chile (Con- cepcion Methane Seep area or CMSA, Sellanes & Krylova, 2005) has been reported, and sampling has brought to light many new bathyal species. Some of the associated chemosymbiotic bivalves found (e.g., Ca- lyptogena, Lucinoma and Thyasira have recently been described (Holmes et al., 2005; Oliver & Sellanes, 2005; Sellanes & Krylova, 2005). New species of gastropods have been named (e.g., Trophon concepcionensis, Houart and Sellanes, 2006; Otukaia crustulum and Margarites huloti, Vilvens and Sellanes, 2006). In the present article we add to this list a new buccinid species from this seep area as well as two new species from the north to central Chile margin. The goal of the present paper is thus to contribute to Page 98 ihe WVelicer, Vols 0sINow the knowledge of the family Buccinidae from north to south-central Chile and to continue the effort of describing the malacofauna of the CMSA ABBREVIATIONS AGT Agassiz trawl CMSA Concepcion Methane Seep Area JS collection of Javier Sellanes, Chile KF collection of Koen Fraussen, Belgium MNHN Muséum National d’Histoire Naturelle, Paris, France MNHNCL Museo Nacional de Historia Natural, Santiago, Chile. lv live collected specimen dd empty shell SYSTEMATICS Class: Gastropoda Cuvier, 1797 Order: Neogastropoda Wenz, 1938 Subfamily: Buccinoidea Rafinesque, 1815 Family: Buccinidae Rafinesque, 1815 Genus: Aeneator Finlay, 1927:414. Type species. Verconella marshalli Murdoch, 1924 (by original designation). Fossil, Tertiary, New Zea- land. Definition. The genus Aeneator is present mainly in the West Pacific, with the geographical center situated around New Zealand, and with an important fauna off southern West America. For an overview of the genus off New Zealand, we refer to Powell (1979:201—203). Here we follow the opinion of McLean & Andrade (1982:12-13) and use Aeneator in a broad sense, without subgeneric splitting for the Chilean species. Three species were previously known from Chilean waters: Aeneator fontainei (d’ Orbigny, 1839), Aeneator (Ellicea) loisae Rehder, 1971 and Aeneator castillai McLean & Andrade, 1982. Aeneator prognaviter new species (Figures 1—2, 12-15) Type material. Holotype(MNHNCL-5863) (32.2 mm), Chile, off Antofagasta 22°51’99 S, 70°29’40 W, in 318 m, lv. Paratype 1 (KF-5178) (26.0 mm), same locality as holotype, lv; paratype 2 (MNHNCL-5864) (29.2 mm), same locality as holotype, dd. Type locality. Chile, north of Antofagasta, Chilean upper continental slope, in 318 m. Range and habitat. Only known from the type material. Description. Shell small for genus (up to 32.2 mm), thick, solid, snow white. Shape broad with moderately high spire, whorls convex, slightly angulate, suture deep. Upper whorls and protoconch eroded, about 4 1/2 teleoconch whorls remaining of which only 3 1/2 with sculpture intact. Spire whorls with 8 or 9 broad spiral cords with rather sharp top, interspaces broad, of equal size. Body whorl with 20—24 spiral cords, occasionally alternating fine and sharp. Spire whorls with 17-19 pronounced, slightly curved axial ribs, interspaces deep, broad. Body whorl with 22 such axial cords, gradually becoming weaker towards outer lip. Aperture oval, columella smooth, slightly curved, outer lip thin, simple, edge sharp. Siphonal canal short, broad, open. Operculum small, thin, transparent, yellowish brown, elongate, nucleus terminal, tip sharp. Comparison. Aeneator prognaviter sp. nov. is char- acterized by broad shape with angular whorls, curved axial ribs and short siphonal canal. Aeneator loisae Rehder, 1971 and A. castillai McLean & Andrade, 1982 both differ in having a higher number of spiral cords in combination with a lower number of axial ribs, spiral cords with a convex top (instead of sharp) and usually a higher number of secondary spiral cords (instead of alternating fine and strong), axial ribs which are straight (instead of bent and curved) and a larger adult size. Aeneator recens (Dell, 1951) from New Zealand is somewhat similar in shape, axial sculpture, size and colour but differs by having narrower spiral cords with broader interspaces, a longer siphonal canal and a curved operculum. Etymology. Aeneator prognaviter sp. nov. is named after the Latin expression “prognaviter,’ meaning “clearly” and “brief and to the point” or “short but sweet” (as adverbum), or meaning “also” (as sub- stantivum), which refers to the shell which is clearly an Aeneator. It also refers to the small size (short or brief) but still an Aeneator (to the point). Aeneator portentosus new species (Figures 3-4, 7-11) Type material. Holotype (MNHNCL-5865) (44.9 mm, siphonal canal broken), Chile, continental slope off Iquique, 21°19’ S, 70°26’ W, in 605 m, dd. Paratype (KF-0338) (45.5 mm), off Coquimbo, 800 m deep, trawled by fisherman. Type locality. Chile, off Iquique, 21°19’ S, 70°26’ W, in 605 m. Range and habitat. Only known from the type material. Description. Shell medium (up to 45.5 mm), thin but solid, snow white. Shape elongate with high spire. Whorls angulate, upper spire whorls rather pagodoid. Spiral sculpture dominant. K. Fraussen & J. Sellanes, 2007 Page 99 Figures 1-2. Aeneator prognaviter sp. nov., holotype, 32.2 mm, Chile, off Antofagasta 22°51'99 S, 70°29'40 W, 318 m, MNHNCL- 5863. Figures 3-4. Aeneator portentosus sp. nov., holotype, 44.9 mm, Chile, off Iquique 21°19S, 70°26W, 605 m,, MNHNCL-5865. Figures 5-6. Kryptos explorator sp. nov., holotype, 29.6 mm, Chile, northwest of the Bay of Concepcion 36°20'97 S, 73°44'86 W, 850 m, MNHNCL-5866. Page 100 The Veliger, Vol. 50, Figures 7-11. Aeneator portentosus sp. noyv., paratype , 45.5 mm, Chile, off Coquimbo, 800 m, KF-0338. No. N K. Fraussen & J. Sellanes, 2007 Page 101 Figures 12-13. Aeneator prognaviter sp. nov., paratype 1, 26.0 mm, Chile, off Antofagasta 22°51’99 S, 70°29'40 W, 318 m, KF- 5178. Figures 14-15. Aeneator prognaviter sp. nov., paratype 2, 29.2 mm, Chile, off Antofagasta 22°51’99 S, 70°29'40 W, 318 m, MNHNCL-5864. Figures 16-20. Kryptos explorator sp. nov., 16-19. paratype 3, 28.9 mm, Chile, northwest of the Bay of Concepcion 36°22'68 S, 73°42'46 W, 708-709 m, KF-5180. 20. operculum of holotype, 6.6 mm. Page 102 Upper whorls and protoconch eroded. All whorls with 6 or 7 sharp spiral cords, subsutural cord weak, gradually stronger along subsutural slope, pronounced on periphery, forming a carina. Interspac- es broad, bottom weakly concave. Body whorl with about 20 spiral cords, 3 or 4 weak ones on subsutural slope, 2 or 3 strong ones on periphery, gradually becoming slightly weaker towards siphonal canal. Upper spire whorls eroded but numerous fine axial ribs still traceable, more pronounced on periphery and on top of spiral cords. Axial ribs gradually weaker towards penultimate whorl. Body whorl smooth. All whorls covered by fine incremental lines. Aperture round, columella smooth, slightly curved, outer lip thin, simple, edge sharp. Siphonal canal moderately short, broad, open, slightly bent. Periostracum (paratype 1) thick, ornamented with a dense sculpture of fine, sharp axial lamellae, running from suture to suture, forming sharp spines or hairs on transition with spiral sculpture. Operculum small, thin, corneous, dark brown, elongate, nucleus terminal, tip sharp. Comparison. Aeneator portentosus sp. nov. is char- acterized by the rather pagodoid shape, the pro- nounced spiral sculpture and the densely sculptured periostracum. The generic placement is based on conchological characteristics and on the shape of the operculum. All Aeneator species known from Chile differ by having more convex whorls, a slightly longer siphonal canal, narrower spiral interspaces and a smoother periostracum. Etymology. Aeneator portentosus sp. nov. is derived from the Latin expression portentosus (adjective), meaning “‘wonderful,” which refers to the graceful shape and excellent sculpture. Genus Kryptos Jeffreys in Dautzenberg & Fischer, 1896 Type species. Kryptos elegans Jeffreys in Dautzen- berg & Fischer, 1896 (type locality: ““bathyal, W. of Spain” designated by Bouchet & Warén, 1985:196), by monotypy, a junior synonym of Pleurotomella koehleri Locard, 1896. Transferred to Buccinidae by Bouchet & Warén (1985:195), based on morphology of the radula. Range, until the present paper, restricted to the Atlantic Ocean, the two known species being K. koehleri (Locard, 1896) (= Kryptos elegans Jeffreys in Dautzenberg & Fischer, 1896; Pleurotomella atlantica Locard, 1897 and Pleurotomella demulcata Locard, 1897) from the N. E. Atlantic and K. tholoides (Watson, 1882) from the S. W. Atlantic (off Brazil). Remarks. Kryptos is characterized by a multispiral, rather big protoconch with a slightly flattened tip, The Veliger, Vol. 50, No. 2 sculptured towards transition to the teleoconch (Fig- ure 28), a smooth, narrow subsutural band, a slightly broader interspaces on the periphery, carinated whorls (type species) or sculptured with some sharp keels (K. tholoides). Bouchet & Warén (1985:196) noted that K. koehleri lack eyes. Americominella Klappenbach & Ureta, 1972 (type species: Americominella duartei Klappenbach & Ureta, 1972) from the Patagonian continental shelf is similar in protoconch morphology and sculpture but differs by the radula, which has a tricuspid central tooth with broad base. Kapala Ponder, 1982 (type species: Kapala kengra- hami Ponder, 1982) from Australia has a radula with an identical central tooth but which differs by having the lateral teeth with 1 large outer cusp and more than 5 small inner cusps. Antarctoneptunea Dell, 1972 (type species: Fusitriton aurora Hedley, 1916) is similar in shape but differs in having a large papilliform protoconch (similar to Aeneator) and a radula with tricuspid central tooth. The new species described below is tentatively placed in Kryptos based on similarities in radula, protoconch and spiral sculpture. Kryptos explorator new species (Figures 5—6, 16—25) Type material. Holotype (MNHNCL-5866) (29.6 mm), south-central Chile, R/V Vidal Gormaz (SeepOx cruise, AGT 6-7, 09/02/2006), CMSA, northwest of the Bay of Concepcion 36°20'97 S, 73°44’86 W, 850 m, lv. Paratype 1 (MNHNCL-5867) (29.4 mm), same locality as holotype, lv; paratype 2 (MNHNCL-5868) (29.3 mm), same locality as holotype, lv; paratypes 3 & 4 (KF-5180—-5181) south-central Chile, R/V Vidal Gormaz (VG-04 Cruise, AGT 10, 10/14/2004), CMSA, northwest of the Bay of Concepcion, 36°22'68 S, 73°42'46 W, 708-709 m; paratype 5 (MNHN-9961) same locality of paratypes 3 & 4. Type locality. South-central Chile, R/V Vidal Gor- maz (SeepOx Cruise, AGT 6-7, 09/02/2006), CMSA, northwest of the Bay of Concepcion, 36°20'97 S, 73°44'86 W, 850 m. Range and habitat. Only known from the type material. All the specimens of K. explorator sp. nov., so far collected have been associated with fauna typical of methane seeps (vesicomyid, solemyid, lucinid and thyasirid bivalves). However, the scarce knowledge of the bathyal SE Pacific malacofauna still prevents us from establishing if this new species lives in an obligate association with seep environments. Description. Shell small (up to 29.6 mm), thin but solid, semi-transparent, white. Shape fusiform with slender spire. Protoconch multispiral, consisting of about 2 1/4 K. Fraussen & J. Sellanes, 2007 Page 103 Figures 21-25. Kryptos explorator sp. nov., holotype, Chile, northwest of the Bay of Concepcion 36°20'97 S, 73°44’86 W, 850 m, MNHNCL-5866. 21. frontal view of removed animal. 22. left side view. 23. close up of the head showing the remarkable eyes. 24. radula, scalebar: 100 micrometer. 25. radula, scalebar: 10 micrometer. Page 104 The Veliger, Vol. 50, No. 2 Figure 26. Figures 27-35. Krypthos tholoides (Watson, 1882), holotype, 16.1 mm, BMNH, after Bouchet & Warén, 1986, fig. 96. Kryptos koehleri Locard, 1896. 27. 21.8 mm, Gulf of Biscay, BIOGAS CP25, 44°05'N, 04°17'W, 1894 m, after Bouchet & Warén, 1985, fig. 511. 28-29. 11.8 mm, off Portugal, after Bouchet & Warén, 1985, fig. 512. 30-31. holotype of Pleurotomella elegans Jeffreys in Dautzenberg & Fischer, 1896, 12.0 mm, MNHN-6422. 32-33. holotype of Pleurotomella atlantica Locard, 1897, 16.5 mm, MNHN-6647. 34-35. holotype of Pleurotomella demulcata Locard, 1897, 13.2 mm, MNHN-6645. K. Fraussen & J. Sellanes, 2007 whorls, about 1.6 mm in diameter, tip flattened, last whorl rather big, convex, ornamented with a reticulate sculpture of 7 or 8 fine spiral cords and numerous fine axial lamellae. Sculpture appearing as small holes when slightly eroded (first protoconch whorl). Transition to teleoconch indistinct. Teleoconch whorls up to 7 in number, convex, adapical part slightly flattened, accentuating a conical shape. Suture distinct. First whorl with 7 spiral cords, at first smooth and weak, gradually becoming stronger and more convex, with deep interspaces of equal width. Second whorl with 8 sharp, narrow spiral cords, interspaces twice as broad. Spiral cords suddenly broader and weaker, but occasionally still sharp, with variable interspaces, usually narrow. Third whorl with 12 spiral cords of mixed strength. Body whorl adapically rather smooth, with numerous weak or obscure spiral cords; base strongly sculptured with about 9 strong spiral cords. Siphonal canal rather smooth with about 15 weak spiral cords. First teleoconch whorl with fine axial riblets at beginning, gradually becoming stronger, waving on top of spiral cords, second whorl with pronounced, sharp, narrow axial ribs, slightly weaker near sutures. Second whorl with 13, third whorl with 14 such ribs. Penultimate and body whorl with 17 axial ribs on adapical half of body whorl, base smooth. All whorls covered with fine, slightly curved incremental lines. Aperture round, columella gently curved, callus thin, smooth. Outer lip thin, sharp, laterally curved accord- ing to incremental lines. Siphonal canal narrow, rather short, open. Operculum corneous, thin, pale brownish, elongate, nucleus terminal, tip sharp. Periostracum yellowish to pale brown, thin, smooth, well adherent. Radula (Figures 24-25) typical for genus: central tooth rather rectangular with concave base and | short cusp, lateral.teeth tricuspid with large outer cusp and small middle cusp. Animal (Figures 21—23) pale yellowish, with 2 short but broad tentacles and black, rather big eyes. Comparison. Kryptos explorator sp. nov. is charac- terized by having a fusiform shape, a multispiral, rather big protoconch with a slightly flattened tip and a reticulate sculpture near the transition to teleoconch, a smooth, narrow subsutural interspace between suture and shoulder and slightly broader, smooth interspaces on the periphery. K. koehleri (Locard, 1896) (Figures 27-35) differs by having a broad shape with strongly angulate whorls and by lacking eyes. K. tholoides (Watson, 1882) (Figure 26) differs by having 2 strong spiral folds, broader interspaces and a glossy surface. Page 105 Etymology. Kryptos explorator sp. nov. is named after the Latin expression explorator (subst., m) meaning “‘a scout” or “the one who search out,” which refers to the range (the Pacific, new for the genus and far from the Atlantic) where this new species is found. It also refers to the presence of eyes (to explore the new habitat visually) which are absent in the type species (K. koehleri). Acknowledgments. We are thankful to Kevin Monsecour (Belgium) for digital images and to David Monsecour (Belgium) for reading and correcting the English text. We also thank the captain and crew of AGOR Vidal Gormaz of the Chilean Navy for support at sea and Guillermo Guzman from Universidad Arturo Part, Iquique who obtained the holotype of A. portentosus. This work was partially funded by Fondecyt project No. 1061217 to J.S. and the research Direction and COPAS center of the University of Concepcion, Fondecyt project No. 1061214 to Praxedes Munoz; NOAA Ocean Exploration Program via SCRIPPS Institution of Oceanography, contract nr. NOAA NAI7RJ1231, and the Office of Naval Research of the US Navy provided extra funding for ship time. REFERENCES ANDRADE, H. 1986. Observaciones biologicas sobre inverteb- rados demersales de la zona central de Chile. In: P. 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Journal of the Linnean Society of London 16:247-254. THE VELIGER oy The Veliger 50(2):107—119 (June 20, 2008) © CMS, Inc., 2007 Redescription of the Deep-sea Wood Borer Neoxylophaga teramachii Taki & Habe, 1950 and its Assignment to the Genus Xy/oredo (Bivalvia: Myoida: Pholadoidea) with Comments on Fossil Pholadoidae TAKUMA HAGA Department of Biological Science, Graduate School of Sciences, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan TOMOKI KASE Department of Geology and Paleontology, National Museum of Nature and Science [formerly National Science Museum, Tokyo], 3-23-1 Hyakunin-cho, Shinjuku-ku, Tokyo 169-0073, Japan (e-mail: haga@kahaku.go.jp) Abstract. Examination of the type specimens of the Japanese wood-borer Neoxylophaga teramachii Taki & Habe, 1950 and additional live and dead intact specimens has revealed that the species should be reallocated to the genus Xy/oredo Turner, 1972, the third genus of Xylophagainae, because it has: 1) a teredo-like calcareous tube, 2) a greatly reduced calcareous mesoplax, and 3) extended inhalant and exhalant siphonal canals, all features typical of Xyloredo. Mesoxylophaga Habe, 1977, established for N. taramachii, is therefore regarded as the junior synonym of Xyloredo. Since the present species is extremely rare and its soft parts never studied, a detailed redescription is given for shell, mesoplax, calcareous tube and soft parts. Two unique organs considered to function in self-fertilization, the accessory genital organ and vesicula seminalis, are observed for the first time for Xyloredo. The associated calcareous tube shows unique microstructure referred to isolated crystal morphotypes. The mineralogy of the calcareous tube consists entirely of aragonite, and the general morphology of the tube is characterized by remarkably strong, regular growth lines in its surface. These features in the calcareous tube provide useful criteria for identification of the trace fossil Teredolites in the Mesozoic and Cenozoic. INTRODUCTION The Xylophagainae is a subfamily of Pholadidae and, according to Turner (2002), composed of three genera, Xylophaga Turton, 1822, Xylopholas Turner, 1972a, and Xyloredo Turner, 1972b. All species of this subfamily are obligate borers in sunken woods, mostly in deep seas: Knudsen (1961) described 17 species of Xylophaga in the collection obtained during the Galathea Deep Sea Expedition from 1950 to 1952. Subsequently, Turner (2002) revised previously report- ed species and described seven additional new species. Both authors demonstrated that the soft parts, such as siphons, mesoplax and accompanying external mor- phology (e.g., ‘chimney’ and ‘tube’ seen in their burrows) are important characters for the systematics of this subfamily; shell characters are generally not useful because they share almost homogeneous, simple, spherical, and Teredo-like valves. Therefore, detailed examination of intact live animals is indispensable for the systematics of this bivalve group. However, intact live animals are quite difficult to obtain due to their deep-sea occurrence and the fragile nature of the shells. Japanese authors proposed four subgenera within the genus Xylophaga all diagnosed by shell characters alone. These are Protoxylophaga Taki & Habe, 1945; Neox- ylophaga Taki & Habe, 1945; Metaxylophaga Taki & Habe, 1945; and Mesoxylophaga Habe, 1977. Turner (1969, 2002), Hoagland & Turner (1981), and Hoagland (1983) synonymized all the subgenera with Xylophaga. However, such taxonomic treatments need to be con- firmed on the basis of detailed examination of soft parts. Neoxylophaga teramachii Taki & Habe (1950) is a case of such examples. It was described on the basis of specimens from Tosa Bay, Kochi Prefecture in western Japan. Subsequently, Habe (1977) established the monotypic new subgenus Mesoxylophaga under the genus Neoxylophaga with N. teramachii as the type species. Kuroda & Habe (1981) later ranked Mesox- ylophaga at the genus level, and this taxonomic treatment has been followed by subsequent Japanese authors (e.g., Higo & Goto, 1993; Higo et al., 1999). Okutani (2000) considered Mesoxylophaga as a subge- nus of Xy/ophaga without any discussion. N. teramachii is very rare, and Pailleret et al. (2007: p. 237, fig. 3) documented for the first time since the description by Page 108 the Veliger, Wol-s03NowZ Figure 1. Type specimens of Xyvloredo teramachii deposited in the Toba Aquarium labeled as TA-7097. A—-B. Holotype. C—F. Paratype #1. G—I. Paratype #2. Scale bars = 5 mm. Taki & Habe (1950) a live animal of this species from a deep-sea bottom off Vanuatu. This paper reexamines the type specimens of N. teramachii deposited in the Toba Aquarium, Toba City, Mie Prefecture, Japan, as well as describes in detail the shell, mesoplax, accompanied calcareous tube, and soft parts of N. teramachii on the basis of additional live and dead specimens obtained by the first author from his recent extensive field sampling. We show that this species can be reallocated to Xyloredo, another genus of Xylophagainae, and that Mesoxylo- phaga is a junior synonym of Xyloredo instead of a ubgenus of Xylophaga. In addition, this paper first ‘uments the mineralogy and microstructure of the careous tube of the genus Xy/oredo. Institutional Abbreviations: NSMT—National Museum of Nature and Science, Tokyo, Japan (formerly National Science Museum, Tokyo); OKCAB—Okayama Univer- sity, Conservation of Aquatic Biodiversity, Okayama, Japan; TA—Toba Aquarium, Mie, Japan. MATERIALS AND METHODS Holotype: Left valve (11.62 mm in height, 12.73 mm in length, TA-7097: Figure 1A—B), possibly taken alive. Taki & Habe (1950) described the present species based on a single conjoined specimen and illustrated it with a freehand figure. The ‘holotype’ deposited in the Toba Aquarium with the registration number TA-7097 consists only of a left valve, although it was ‘conjoined’ T. Haga & T. Kase, 2007 in the original description, of which Higo et al. (2001) were aware. Its dimensions do not match well with those given in the original description. However, this ‘holotype’ specimen matches the figure and retains part of the dried-up posterior adductor muscle within the valve, suggesting that it was originally a conjoined valve. Shell dimensions sometimes differ because of varying measuring methologies. We therefore regard TA-7097 as the holotype. Paratypes: Two conjoined shells (Paratype #1, 11.90 mm in height, 11.15 mm in length: Figure 1C— F; Paratype #2, 8.93 mm in height, 9.91 mm in length: Figure 1G—I), possibly taken alive from the type locality. Although Taki & Habe (1950) did not mention paratypes in the original description, two conjoined specimens were labeled and preserved under the registration number of TA-7097, the same as the holotype (Figure 2). The type specimens deposited in the Toba Aquarium are briefly labeled and holotypes and paratypes are distinguished with red and blue labels, respectively (M. Isowa, personal communica- tion). Although the other two specimens are not marked with blue labels, they are regarded as paratypes. Other material: JAPAN—Five empty shells, one individual with decayed animal and fragments of tubes inside a sunken wood trunk recovered by a commercial trawler at 200 m deep off Tokai, Ibaraki Prefecture, April 29, 2004, leg. T. H. (NSMT- Mo76705); 50 empty shells, two live individuals and nine almost intact tubes, inside a sunken wood trunk recovered by a commercial trawler at a depth of 125 m off Tokai, Ibaraki Prefecture, June 3, 2004, Jeg. T. H. (NSMT-Mo76706; OKCAB M15894); 12 empty shells and fragments of tubes, inside a sunken wood trunk recovered by a commercial trawler at a depth of 500 m off Hitachi, Ibaraki Prefecture, March 10, 2007, leg. T. H. (NSMT-Mo76707); three empty shells and four tubes, inside a sunken wood trunk, 250-300 m deep off Atsumi Peninsula, Aichi Prefec- ture, February 1999, /eg. S. Kimura (NSMT- Mo76708). WANUATU—two intact specimens; BOAO, from a depth of 560-580 m between Malekula and Epi Island, Vanuatu, inside a sunken wood trunk identified either as Leucaena or as Serianthes, Novem- ber, 2004, R/V Alis (NSMT-Mo73806, 73807) (Pailleret et al., 2007). Methods: Since the outer morphology of soft body in situ provides characters indispensable for systematics, we exposed the animals by breaking the wood with special care, and then observed and photographed the specimens. We used one of two live individuals (originally prepared for molecular analysis with 99% ethanol, following Ueshima, 2002) recovered from off Page 109 Figure 2. Type specimens of Xyv/oredo teramachii with the attached label in the Toba Aquarium, registered as TA-7097. A. The label. Japanese texts mean Japanese name (teramachi- kukui-gai = teramachi xylophagain clam), the family (kikui-gai-ka = Xylophagaidae), and the locality (Tosa 100 fms. = Kochi Prefecture, —100 fathoms deep) from top to bottom, respectively. B. Type specimens. Arrowhead indicates the holotype. Note a boxed seal attaching the holotype. Tokai, Ibaraki, Japan in June, 2004 for gross anatomy. This material was strongly dehydrated, so that we employed a method improved from Fukuda & Ekawa (1997): (1) the whole animal was immersed in 5% HCl solution for 30 min to rehydrate it sufficiently for dissection, (2) rinsed with tap water for 10 min, and then (3) dissected in 70% ethanol under a binocular microscope. As the dried periostracal sheath was strongly contracted, it was photographed by immersing in 70% ethanol. Scanning electron microscopy (SEM) micrographs were produced on a JOEL-T330A scan- ning electric microscope after the preparation of material with the standard method: material was cleaned and rinsed with distilled water, hydrated with pure ethanol, dried, mounted on the stages, and then coated with gold. For prodissoconch observation, the shelled structure covering the umbo was removed manually to expose the prodissoconch prior to the preparation. X-ray diffraction analyses were conducted for mineralogical determination of the shell and calcareous tubes. All the specimens used in this study are housed at NSMT (NSMT-Mo 76705-76708). We followed Purchon (1941) and Turner (2002) for the terminology of anatomical and conchological characters, respectively, except for ‘dorsal portion of Page 110 The Veliger, Vol. 50, No. 2 aca Figure 3. Internal left valve of Xyloredo teramachii. Abbreviations: aa = anterior adductor scar, aca = accessory anterior adductor scar, cp = chondrophore, dpa = dorsal portion of posterior adductor scar, pa = posterior adductor scar, pl = pallial line, ppr = posterior pedal retractor scar, sf = shelf, ub = umbo, ubr = umbonal-ventral ridge, ur = umbonal reflection, vc = ventral condyle, vpa = ventral portion of posterior adductor scar. Scale bar = 5 mm. posterior adductor,’ ‘ventral portion of posterior adductor’ and ‘accessory anterior adductor,’ which are newly introduced herein. SYSTEMATICS Superfamily Pholadoidea Lamarck, 1809 Family Pholadidae Lamarck, 1809 Subfamily Xylophagainae Purchon, 1941 Genus Xyloredo Turner, 1972b Xyloredo Turner, 1972b, p. 3. Type species: Xyloredo nooi Turner, 1972b, by original designation. Mesoxylophaga Habe, 1977, p. 295. Type species: Neoxylophaga teramachii Taki & Habe, 1950. Remarks: Turner (1972b) established the genus Xylor- edo by distinguishing it from all the other xylophagines in having 1) a long, teredinid-like burrow with a calcareous lining, 2) a thin periostracal border of the tube, and 3) extended inhalant and exhalant canals. Xyloredo superficially resembles genera of the Teredi- nidae, but its anatomical features indicate placement in Xylophagainae; these features are: 1) the U-shaped wood-storing caecum, 2) the internal visceral mass completely covered by the shell, 3) the presence of mesoplax, and 4) the absence of apophysis and pallet (Turner, 1972b, 2002). Aside from the type species, Turner (1972b, 2002) included Xy/oredo ingolfia Turner, 1972b and Xyloredo naceli Turner 1972b in this genus. We here include Neoxylophaga teramachii Taki & Habe, 1950 as the fourth species of this genus. Therefore, Mesoxylophaga is a junior synonym of Xy/loredo. T. Haga & T. Kase, 2007 Figure 4. Xyloredo teramachii, enlarged view of sagittal section of the anterior slope. NSMT-Mo76706. Prismatic layer and crossed lamellar layer are indicated as A and B, respectively. Scale bar = 100 um. Xyloredo teramachii (Taki & Habe, 1950) [new combination] (Figures 1-11) Neoxylophaga teramachii Taki & Habe, 1950, p. 46, fig. 3; Kuroda et al., 1971, p. 715 (Japanese text), 471 Page 111 (English text), pl. 121, fig. 11; non Okutani, 1968, pasa pleZatieg Ss Neoxylophaga (Mesoxylophaga) teramachii. Habe, 1977, p. 295. Xylophaga teramachii. Hoagland & Turner, 1981, p. 44; Hoagland, 1983, p. 7; Turner, 2002, p. 227 (written as Xylophaga teramachi [sic]); Pailleret et al., 2007, p. 236, fig. 3. Mesoxylophaga teramachii. Kuroda & Habe, 1981, p. 179, pl. 7, fig. 6; Higo & Goto, 1993, p. 670; Higo et al., 1999, p. 520; Higo et al., 2001, p. 181. Xylophaga (Mesoxylophaga) teramachii. Okutani, 2000, p. 1031, pl. 513, fig. 3. Shell: The valve is globose, up to 16.5 mm in height, 17.9 mm in length in the largest specimen, and has a shape typical of the subfamily, consisting of an anterior slope, disc and posterior slope (Figures 1—2). The valve surface is originally pearly white in color and glossy, and covered with a thin, dark golden brown perios- tracum over the whole surface (Figures 1, 8A). Finely denticulated bunches are regularly distributed over the anterior slope from the umbonal-ventral sulcus for- wards, and fine growth lines extend over the disc and posterior slope. The posterior slope is widely reflected dorsally and its ventral midline, where the shelf terminates with the posterior slope and forms a blunt angulation (Figure 3). Since the margin of posterior Figure 5. Prodissoconch of Xyloredo teramachii. NSMT-Mo76706. Arrowheads indicate the boundaries among prodissoconch I, II, and dissoconch. Abbreviations: dc = dissoconch, pdI = prodissonconch I, pdII = prodissoconch IH. Scale bar = 100 um. Page 112 The Veliger, Vol. 50, No. 2 Figure 6. Mesoplax of Xyloredo teramachii. NSMT-Mo76706. A-B. Mesoplax in situ attached to the extended periostracum at the umbonal reflection indicated by arrowheads. C—D. Enlarged ventral view of the mesoplax. Abbreviations: cp = chondrophores, rl = rod-shaped ligament. Scale bars = A—B: 2 mm; C: 500 um; D: 20 um. slope reflects laterally, the posterior end opens widely and large siphonal canals extend from it (Figure 6A). The umbonal-ventral sulcus (Figure 1A, C, G) is wide and concave without the crenate varix. The umbonal reflection largely develops and reflects dorsally, so that it is ear-shaped in anterior view (Figures 1E, 6B) The internal valve surface is smooth, also pearly white and glossy. The shelf largely develops at the middle of the posterior slope and is impressed deeply to form a bump similar to that in teredinids (sf: Figure 3). The umbonal-ventral ridge (ubr: Figure 3) is wide, laterally depressed, irregularly marked with rough lines, descends from the umbo along the middle of the disc, and forms the large spherical ventral condyle (vc: Figure 3). In the left valve, the large, cuneiform and flat chondrophore is prominent beneath the umbo (cp: Figure 3), but in the right valve, the brown- colored, rod-shaped ligament protrudes laterally (Fig- ure 6B). The umbo strongly curves antero-ventrally (ub: Figure 3), and the prodissoconch is completely enclosed in the anterior reflection. The posterior muscular scar (pa: Figure 3) is large, deeply impressed with many lines, and divided into two areas: one with numerous irregular narrow lines that spread inward along the dorsal part of posterior slope (dorsal portion of posterior muscular scar: dpa, in Figure 3), and the other with a few wide lines that obliquely spread ventro-internally and are chevron-shaped ventrally (ventral portion of posterior muscular scar: vpa, in Figure 3). The pallial line (pl: Figure 3) is narrow and located along the valve margin, and is rather obscure near the ventral condyle. The accessory anterior adductor muscle scars (aca: Figure 3) and anterior adductor scars (aa: Figure 3) are rather obscure and T. Haga & T. Kase, 2007 Page 113 Figure 7. Calcareous tubes of Xy/oredo teramachii. NSMT-Mo76706. A—B. Posterior end of the tube. Arrrowhead, dotted line, and box indicate the enlarged views of D-H. C. posterior tip of the tube. Arrowheads indicate the lateral ‘blades’. D. Enlarged view of periostracum which externally covers the tube. E. Enlarged view of sagittal section. F. Enlarged view of the outer surface. G—H. Enlarged view of the inner surface. Scale bars = A—B: 10 mm; D, F: 3 um; E, G: 10 um; H: 2 um. Page 114 The VeliseraV oly 505 iNowZ Figure 8. Periostracal sheath of Xvloredo teramachii which covers the siphonal canals. NSMT-Mo76706. A. Intact specimen in situ with the complete periostracal sheath fully extended. B—C. Enlarged view of microscopic pores, indicated by arrowheads. Scale bars = A: 20 mm; B: 100 um; C: 10 um. Figure 9. A—C. Animal in situ of Xyloredo teramachii. NSMT-Mo76706. Arrowhead indicates the boundary of the calcareous tube and the periostracal sheath. B. A sunken wood trunk in situ bored by Xyloredo teramachii. Abbreviation: ct = calcareous tube. Scale bars: A-—C 5mm; D = 20 mm. T. Haga & T. Kase, 2007 Page 115 Jaye i 2 - — erm = Figure 10. Accessory genital organ of Xyloredo teramachii. A. Posterior view. B. Anterior view. C. Lateral view. Abbreviations: ago = accessory genital organ, an = anus, dpa = dorsal portion of posterior adductor, rt = rectum, vpa = ventral portion of posterior adductor. Scale bar = | mm. fuse with the pallial line; the former hes at the anterior margin of the anterior slope, and the latter are small and are located in the middle of the umbonal reflection. The posterior pedal retractor scar (ppr: Figure 3) is large, located close to the ventral portion of the posterior adductor scar, and positioned in the middle of the shelf. The anterior pedal retractor scar is situated beneath the anterior reflection. The ventral adductor scars and siphonal retractor scars are absent. The valve is entirely of aragonite and consists of an outer, seemingly irregular simple prismatic layer, and an inner crossed-lamellar layer (Figure 4). Very thin prismatic sublayers, possibly of mycostracal prisms, are occasionally inserted into the inner layer. This sublayer is also observed underneath the inner layer where the muscles adhere. The outer prismatic layer becomes thicker and forms denticles particularly in the anterior slope, and the crossed lamellar layer is prominent particularly in the strongly curved regions such as the umbo and the shelf. The umbonal-ventral ridge consists only of simple prisms. Prodissoconch: The prodissoconch (Figure 5) is typical of planktotrophic development and composed of ca. 80 um-long prodissoconch I (pdI: Figure 5) and ca. 205 um-long prodissonconch II (pdII: Figure 5). It is totally concealed within the umbo due to the anterior reflection and subsequently developed dissoconch in full-grown individuals. The surface is rather rough in prodissoconch I, while it is smooth and marked with regularly spaced concentric growth lines in prodisso- conch II. The boundary between prodissoconch I and II is clearly marked with thin crenations (Figure 5, arrowheads). Mesoplax: The mesoplax (Figure 6) is paired, tiny, long, subquadrangular (widening posteriorly), and slightly calcified. It is situated beneath the posteriorly ascended periostracum that covers the large and oval anterior incision latero-anteriorly (Figure 6A—B). The mesoplax consists of granular prisms with ca. 20 um- wide subunits (Figure 6C—D). This structure is difficult to observe in live specimens because of their tiny size and the complete coverage with mucous debris (Figure 9B). Burrow and calcareous tube: The long burrow (Fig- ure 9D) is typical of teredinids, but does not produce ‘nodules,’ indicative of the switch-backed drilling behavior observed in teredinids. The approximate posterior two thirds of the burrow is lined with a calcareous tube (Figure 7) that is marked with distinct, regularly spaced growth rings (Figure 7A—B). The external surface of the tube is totally covered with a Page 116 The Veliger, Vol. 50, No. 2 Figure 11. A. Sagittal section of the siphonal canals. B—C. Enlarged view of the siphons. Abbreviations: cv = cavity, ed = epidermis, es = exhalant sihonal canal and siphon, is = inhalant siphonal canal and siphon, po = periostracum, sm = sigmoid mesenchyme. Scale bars = 1 mm. thin, smooth, and golden-glossy periostracum extend- ing from the valve via a periostracal sheath (Fig- ure 7D). The posterior portion of the tube is rather thick, and bears a lateral ‘blade’ internally at the tip in full-grown individuals (Figure 7C). From the posterior tip to the midpoint, the calcareous tube becomes significantly thinner, and is terminated with a dark- brown, strong demarcation (Figure 9A, arrowhead). The periostracal sheath extended from the valve also terminates at this demarcation (Figure 8A). The siphonal canal between the valve and calcareous tube is therefore solely covered with a periostracal sheath. The surface of periostracal sheath is smooth, but with microscopic holes (inside of pore is ca. 50 um in length) all over the surface (Figure 8B—C). These tiny pores are limited in distribution in the extended periostracal sheath. The tube is composed mostly of periostracum in very young individuals; however, calcification of the tube occurs first at the base of the siphon with simple prismatic structure as well as on the internal surface of the full-grown tubes. The calcareous tube is of aragonite, and consists of alternating fine and coarse hexagonal prismatic fibers, oriented either vertically or horizontally (Figure 7G—H) which gradually become smaller in diameter and length towards the outer portion (Figure 7E—F). The vertical prismatic fibers are usually less than 10 um in diameter, and larger and more abundant than the horizontal prismatic fibers. Soft parts: The animal is long as in teredinids, but differs in having a completely internal visceral mass, a mesoplax at the anterior incision, and in the absence of apophysis and pallet (Figure 9A—C). The anatomical features are generally identical to xylophagaines (e.g., Purchon, 1941; Turner, 2002): the posterior adductor muscle is large, divided into two parts, and the ventral part consists of chevron-shape fibers and occupies two thirds of the posterior adductor, while the dorsal part consists of irregularly arranged fibers and descends along the posterior slope. The anterior adductor is small, depressed dorso-ventrally and sinuous in the anterior reflection. The accessory anterior adductor is T. Haga & T. Kase, 2007 small, situated beneath the anterior adductor, and consists of stout muscular fibers. The posterior pedal retractor is large and closely inserted into the midpoint of the posterior adductor. The anterior pedal retractor is attached within the deeper portion of the umbo. The ventral adductor and the siphonal retractor are absent. The foot is large, discoid, and is surrounded by circular muscles around its margin (Figure 9C). The gills are narrow, thick, laterally stout, and consist only of inner demibranchs. The digestive tract, also typical of xylophgaines, has a large U-shaped wood-storing caecum that is con- nected to the stomach on its left side. The intestine ascends along the anterior margin of the posterior adductor, penetrates the heart, then terminates as a simple anus, which is surrounded by the accessory genital organ (Figure 10). The accessory genital organ is well-developed, glandular, free from any adhesion to the posterior adductor, and superficially composed of two components: a blade surrounding the end of the rectum (Figure 10A), and a peduncle whose ventral end protrudes like a proboscis and is situated below the rectum (Figure 10OB—C). A pair of vesicula seminalis, a flattened small lobule visible in pale yellow, is present laterally on the thin suspensory membrane of the cteni- dium close to the posterior end of the pedal retractor. The siphons (Figure 11B) are short, simple, and both tips are usually aligned with the same length; however, the tip of the exhalant siphon appears to be slightly longer in some individuals. The apertures in both siphons are roughly serrated (Figure 11C). The sipho- nal canals are long and are connected to the posterior part of the visceral mass and siphons. The inhalant and exhalant siphonal canals are supported by well-devel- oped sigmoid mesenchymes (sm: Figure 11A). A pair of cavities (cv: Figure 11A), that probably act as a haemocoel, are situated laterally between the siphonal canals, and continuously extends antero-posteriorly from the bases of siphons and visceral mass. Type locality: Tosa Bay, Kochi Prefecture, western Japan, ca. 100 fathoms deep. Distribution: West Pacific along the Japanese mainland from Ibaraki Prefecture to Kochi Prefecture and Vanuatu in the south Pacific. Depth ranges from 125 to 580 m. DISCUSSION Our detailed study on the shells and soft parts shows that N. teramachii can be reallocated to the genus Xyloredo. Xyloredo nooi, X. ingolfia and X. naceli are all less than 10 mm in maximum shell length (Turner, 1972b), while the present species reaches up to ca. 18 mm. In addition to the large shell size, the present species is easily distinguished from the above three Page 117 species in having a laterally reflected and developed posterior slope and dark golden brown periostracum. Okutani (1968: p. 23) identified a specimen from a sunken timber obtained at a depth of 1,510m in Sagami Bay as Neoxylophaga teramachii (but in the figure caption he indicated it as “?Neoxylophage teramachii’ [sic]). We suggest however that this specimen seems to belong to another, yet undescribed species because it differs from the present species in having a large and thick mesoplax, a flatted umbonal- ventral sulcus, and a varix-like crenation in its posterior portion. Habe (1977) allocated Neoxylophaga lobata (Knudsen, 1961) and N. knudseni Okutani, 1975 to his subgenus Mesoxylophaga. These two species do not have the calcareous tube in the burrow and therefore cannot be referred to Xyloredo. Generic positions of these species still remain uncertain until anatomical details are clarified. The present species is the first record of the genus Xyloredo in the West Pacific. Xyloredo nooi, X. ingolfia and X. naceli were reported from the Atlantic, East Pacific, and South Pacific, respectively, from depths of more than 1,500 m, and they were heretofore known only from their type localities (Turner, 1972b, 2002). Hoagland (1983) suggested oviparous development for the above three species, but the present species appears to undergo planktotrophic development judging from the size (ca. 80 um in length) of prodissoconch I (see Jablonski & Lutz, 1980). The wide distribution from Japan to Vanuatu of the present species is likely due to its planktotrophic larval transport. Teredinidae and Xylophagainae have unique repro- ductive strategies. Turner (1968) and Turner & Johnson (1971) stated that Teredinidae and Xylophagainae studied so far exhibit protandrous hermaphroditism, and suggested self-fertilization for Xylophagainae. Purchon (1941) extensively studied the mechanism of self-fertilization in Xylophaga dorsalis (Turton, 1819), and observed spawned sperms deposited in the seminal receptacle via the accessory genital organ that functions to tangle flooded sperms. Hoagland (1983) mentioned that all species of Xy/oredo lack the accessory genital organ. We nevertheless recognized this organ in the present species (Figure 10) and an undetermined species of Xyloredo from Japan. Therefore, our study confirms the presence of the accessory genital organ in Xyloredo. It seems likely that the present species also undergoes self-fertilization because of the presence of vesicula seminalis and an accessory genital organ. However, this conclusion must be confirmed by detailed histological study on individuals with different developmental stages. We show that the calcareous tube of Xv/oredo is composed of hexagonal prismatic fibers, composed of aragonite. The prisms are vertically and horizontally oriented across each other (Figure 7G—H), and its Page 118 The Veliger, Vol. 50, No. 2 Table 1 Mineralogical and morphological features of the calcareous tubes in Xyloredo, Teredina, and Teredinidae. Note that Teredina is extant genus. Family Genus Mineralogy Gross morphology Surface morphology Pholadidae Xyloredo aragonite! long, winded'** maked by regular growth lines’? Teredina aragonite* long, straight?® smooth, gaped at dorsal and ventral?® Teredinidae calcite and/or aragonite*’ long, winded>* smooth?* References: 'this study; 7Turner (1972b); *Turner (2002); *Boggild (1930); °Turner (1969); “Kelly (1988); 7Carter (1980b): *Turner (1966). structure is referable to ‘isolated crystal morphotypes’ as defined by Carter (1980a) and Carter & Clark (1985). Carter (1980a) reported that this microstructure is seldom observed in bivalves, since it has rarely been discussed in detail (e.g. Boggild, 1930). As far as we are aware, the aragonitic isolated crystal morphotypes in Xyloredo reported herein is a characteristic microstruc- ture among the accessory calcareous tubes in bivalves. In Pholadoidea, only wood-borers produce long calcareous tubes: those are the genus Xy/oredo (Turner, 1972b, 2002), fossil genera Teredina Lamarck, 1818 and Turnus Gabb, 1864 (Turner, 1969; Kelly, 1988) in Pholadidae, and all members of the family Teredinidae (Turner, 1966, 1969). Their fossilized burrows were described under the ichnogenus Teredolites Leymerie, 1842, which is characterized by a large club-shaped morphology with a single aperture, and occurs in xylic substrata since the Mesozoic (Leymerie, 1842; Hatai, 1955; Turner, 1966; 1969; Bromley et al., 1984; Plint & Pickerill, 1985; Kelly, 1988). Identification of the trace makers for Teredolites is, however, generally difficult because the burrows usually do not preserve internally embedded body fossils such as valves and/or palletal structure (Plint & Pickerill, 1985). Polychaetes and boring isopods produce similar burrows in xylic substrata, but they never secrete calcareous tubes in their burrows (Gingras et al., 2004). Therefore, Teredolites with the calcareous tubes can be attributed to pholadoidean boring bivalves. We suggest that Teredolites associated with fossilized calcareous tubes can be referred to a specific family or genus within Pholadoidea by analyzing the mineralogy and external tube morphology (Table 1). Namely, Xyloredo is characterized by having a_ winding, aragonitic tube with remarkably strong, regular growth lines on the surface. In Teredina (an odd tube-bearing fossil genus of Pholadidae with morphology convergent to Teredinidae), the tube is mostly straight and aragonitic in composition but its surface is smooth with two gapes at the dorsal and ventral portions in some individuals (Boggild, 1930; Turner, 1969; Kelly, 1988). On the other hand, Teredinidae has a strongly winding, calcitic and/or aragonitic tube with a smooth tube surface (Turner, 1966, 1969; Carter, 1980). Turnus is a poorly known fossil genus originally placed in Teredinidae (Gabb, 1864; see also Turner, 1969) mostly from the Cretaceous. Kelly (1988) reported a lined calcareous Teredolites associated with valves of Turnus kotickensis Kelly, 1988, and tentatively referred the genus to Pholadidae. However, the systematic position of Turnus remains unclear until mineralogical and microstructural information is available. In conclusion, the mineralogy and external morphology of calcareous tube provide useful criteria for identification of Teredolites from the Mesozoic and Cenozoic. Acknowledgments. We thank Messrs. M. Isowa, Y. Yamazaki and M. Furuta (TA) who allowed us to examine the type specimens, Dr. R. Miyawaki (NSMT) for his X-ray diffraction analysis, Dr. M. Zbinden (Pierre and Marie Curie University, Paris) for the access of the samples from Vanuatu, Messrs K. Kurosawa and Y. Gorai (Ibaraki Prefecture) for collection of specimens using their commercial trawlers, Mr. S. Kimura (Mie Prefecture) for providing his specimens, Dr. T. Sasaki (The University Museum, The University of Tokyo) for valuable discussions. We also thank anonymous reviewers and Dr. G. J. Vermeij (University of California, Davis) for helpful comments and suggestions. This study was funded by a grant from the Japan Society of the Promotion of Science (JSPS) (no. 198300) to T. H., and grants from JSPS (no. 18253007) and the National Museum of Nature and Science, Tokyo to Ke. LITERATURE CITED BOGGILD, O. B. 1930. The shell structure of the mollusks. Det Kongelige Danske Videnskabernes Selskabs Skrifter. Naturvidenskabelig og Mathematisk Afdeling 9:231—326; 14 pls. BROMLEY, R. G., S. G. PEMBERTON & R. A. RAHMANTI. 1984. A Cretaceous woodground: the Teredolites ichnofacies. Journal of Paleontology 58(2):488—498. CARTER, J. G. 1980a. Controls of bivalve mineralogy and microstructure. Pp. 69-113 in D. C. Rhoads & R. A. Lutz (eds.), Skeletal Growth of Aquatic Organisms. 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Organization for Economic Co-operation and Development: Paris. UESHIMA, R. 2002. Simple methods for DNA preservation in molluscan specimens. Venus 61(1/2):91—94. (in Japanese with English abstract). The Veliger 50(2):120-128 (June 20, 2008) THE VELIGIR © CMS, Inc., 2007 A Note on Strombus coronatus Defrance, 1827 and Strombus coronatus Roding, 1798 (Mollusca: Gastropoda) MATHIAS HARZHAUSER Naturhistorisches Museum in Wien, Burgring 7 - A-1010 Vienna, Austria (e-mail: mathias.harzhauser@nhm-wien.ac.at) GIJS C. KRONENBERG Milieu Educatie Centrum, Postbus 435, NL-5600 AK Eindhoven, The Netherlands (e-mail: gijs.kronenberg@tiscali.nl) Abstract. Strombus coronatus Defrance, 1827 is considered a nomen protectum, and the cerithiid Strombus coronatus Réding, 1798 is demonstrated to be a nomen oblitum in accordance with the ICZN Article 23.9.1. The Late Miocene to Early Pliocene Strombus coronatus Defrance is shown to belong to Persististrombus Kronenberg & Lee, 2007, and its relation with other Neogene strombids is briefly addressed. A lectotype for Strombus coronatus Defrance is designated. Key Words: Gastropoda, Cerithiidae, Strombidae, Persististrombus, homonymy, Miocene, Pliocene. INTRODUCTION While reviewing literature in the course of our research on strombid taxa from the Miocene Central Paratethys Sea, the nominal taxon Strombus coronatus Defrance, 1827 was frequently encountered. Although the identity and stratigraphic range of this species varies consider- ably in the literature (ranging from Late Oligocene to Late Pliocene) this name is treated as the valid name for a species of the family Strombidae Rafinesque, 1815. Nevertheless, the binomen Strombus coronatus was first introduced by Réding (1798) for an extant Indo-Pacific cerithiid. At that time the generic concept of Strombus differed fundamentally from the modern usage, and an analogous situation exists involving Strombus granula- tus R6ding, 1798 (a cerithiid) and the strombid Strombus granulatus Swainson, 1822 (Kronenberg and Lee, 2005). NOMENCLATURAL STATUS AND HISTORY 1. Strombus coronatus Roding, 1798 When introducing Strombus coronatus, Roding (1798:98, species 1270) referred to Murex aluco Gmelin (1791, sp. 134) and Martini (1777:figs. 1478-1479). Houbrick (1978:104—-105) pointed out that Rédding’s (1798) reference to Martini (1777) involves two species, which are now established as Pseudovertagus aluco (Linnaeus, 1758) [fig. 1478 of Martini (1777)] and Rhinoclavis vertagus (Linnaeus, 1758) [fig. 1479 of Martini (1777)] respectively. Although Houbrick (1978) did not explicitly designate a lectotype for Strombus coronatus Roding, 1798, he synonymized Strombus coronatus Réding, 1798 with Murex aluco Linnaeus, 1758 [= Pseudovertagus aluco (Linnaeus, 1758)] by stating: “I here restrict Strombus coronatus [of Rdding, MH and GCK] to fig. 1478 and place it into the synonymy of Pseudovertagus aluco ...”’ (Houbrick, 1978:104-105). Réding’s S. coronatus soon disappeared from the literature, and the authors know of no 20th century reference to this binomen as a valid species name. 2. Strombus coronatus Defrance, 1827 Defrance (1827) introduced the primary homonym Strombus coronatus, 29 yr after Roding, referring to the illustration of a fossil strombid illustrated in Walch (1768:116, pl. C (38), figs. 1-2). Walch (1768) described the shell as a rare species present in collection cabinets of the time and suggested that it was collected in the Turin region of Italy. Defrance (1827) provided an adequate description of the shell, compared its morphology briefly with the extant Strombus gigas Linnaeus 1758, and remarked that it is frequently found in the Siena region in Italy. He seems to have doubted the Turin origin suggested by Walch (1768) and emphasized that the origin of that specimen was unknown. Indeed, the large-sized species is very common in the Italian Pliocene and was already known to science in the 17th century when Aldrovandi (1648) M. Harzhauser & G. C. Kronenberg, 2007 identified it as Murex albus. After the definition by Defrance (1827), the nominate taxon was cited frequently from Pliocene deposits throughout the Mediterranean and Eastern Atlantic regions. Unfortunately, the species name was also applied to several Miocene and even Oligocene specimens from central and southern Europe. Most notably among others, Hérnes (1856) and Hoernes & Auinger (1884) identified Middle Miocene specimens of what is here provisionally called the Persististrombus lapugyensis- exbonellii group as Strombus coronatus Defrance, 1827. Subsequently, many stout and strongly sculptured fossil strombids with long shoulder-spines have been treated as Strombus coronatus (e.g., Baldi, 1973; Schultz, 1998). 3. Strombus coronatus, Defrance, 1827 — a nomen protectum Herein, we refer to the International Commission for Zoological Nomenclature (ICZN) Article 23.2, which pleas for nomenclatural stability and perpetuation of long-accepted names, and to ICZN Article 23.9.1. To our knowledge, the name Strombus coronatus Roding, 1798 has not been used as a valid name after 1899, which meets the requirements of ICZN Article 23.9.1.1. (“the senior homonym has not been used as a valid name after 1899”). ICZN Article 23.9.1.2. states that “the junior homonym has been used as its presumed valid name in at least 25 works, published by at least 10 authors in the immediately preceding 50 yr and encompassing a span of not less than 10 yr.” Accord- ingly, we document that Strombus coronatus Defrance, 1827 was used between 1956-2006 in 33 papers by 33 authors (see references below), which meets the requirements of Article 29.9.1.2. Thus, we invoke ICZN Article 23.9.1 to make the name Strombus coronatus Defrance, 1827, a nomen protectum and Strombus coronatus Réding, 1798 a nomen oblitum. CURRENT STATUS OF STROMBUS CORONATUS DEFRANCE, 1827 — SYSTEMATICS AND PALEOBIOGEOGRAPHY Recently, Kronenberg & Lee (2007) introduced Persis- tistrombus (type species by original designation: Strom- bus granulatus Swainson, 1822) as a new genus for a group of strombids which experienced its acme in the European Miocene and is now represented by Persis- tistrombus latus (Gmelin, 1791) in the African Eastern Atlantic Province and by P. granulatus (Swainson, 1822) in the Panamic Province. Although this grouping would make Persististrombus seemingly paraphyletic (see the consensus tree present- ed by Latiolais et al. (2006:440)), we here advocate the possibility of a distinct lineage, with hardly any morphological change since the Early Miocene, with two distinct side branches, one leading to Strombus Page 121 (here used in the strict sense, 1.e., represented by the Recent species S. pugilis Linnaeus, 1758 (type species); S. alatus Gmelin, 1792; S. gracilior Sowerby, 1825) and one to Lobatus Iredale, 1921 (type species by mono- typy: Strombus bituberculatus Lamarck, 1822 = Strom- bus raninus Gmelin, 1791). Moreover, the tree present- ed by Latiolais et al. (2006) is a consensus tree, based on 325 bp nuclear histone H3, where indeed P. granulatus plots out as the sister taxon of Strombus s.s. (Latiolais, 2003:fig. 1). These clades are sister to Lobatus for 640 bp mitochrondial COI. Strombus granulatus plots out as sister taxon of Lobatus (Latiolais, 2003:fig. 2) and these two are sister to Strombus 8.8. Persististrombus is characterized by **... moderate size for family, fusiform, shoulder knobs distinct on body whorl, slightly expanded outer lip with sharp, unglazed rim and no extensions, regularly divided callus on columella, anterior canal short, posterior canal or groove absent or obsolete. Protoconch with four to five smooth whorls. Adaxial side of outer lip smooth, plicate, or granulate.”” (Kronenberg and Lee, 2007). Strombus coronatus fits within this definition except for its low, concave spire and the number of protoconch whorls. A preliminary analysis of the Persististrombus lapugyensis-exbonellii group (Harzhauser and Kronen- berg in prep.) reveals that there 1s a gradual change in spire height, i.e., from high-spired specimens in the early Langhian to lower spired specimens in the Serravallian of the Central Paratethys. Therefore we allocate both S. coronatus and the Pliocene to Recent S. /atus Gmelin, 1791 to Persististrombus. As the protoconch in all examined specimens was poorly preserved, the number of whorls may have been slightly higher than the approximately three observed by us (see below). On the other hand, reduction of the number of protoconch whorls may have occurred in the Proto-Mediterranean, which would call for a minor adjustment in the description of Persististrombus as far as the number of protoconch whorls is concerned. Genus Persististrombus Kronenberg and Lee 2007 Numbers in front of references refer to citations which are relevant for ICZN Article 23.9.1.2 (references before 1958 are found in the text above). Persististrombus coronatus (Defrance, 1827) nov. comb. Pl. 1, Figures 1-5, 7-9 Murex albus Aldrovandi, 1648:472, fig. 2. Porphyroides Lancisi, 1771:298, fig. 1. stumpfgestachelte dicklippigte Fligelschnecke Walch, 1768:116, pl. C (38), figs. 1-2. Strombus coronatus Defrance, 1827:124. The Veliger, Vol. 50, No. 2 io we (ooLE FULOCOOL2,... Wg Mt20tt de biaiehs Figures 1-2. 38, figs. 1-2). Figures 3-6. Figures 7-9. Copy of the illustration of the lectotype (designated herein) of Strombus coronatus Defrance, 1827 in Walch (1768, pl. Persististrombus coronatus (Defrance, 1827). 3—5: Specimen 93(1790) present in NMB. 6: Label with NMB specimen. A typical representative of Persististrombus coronatus from the Lower Pliocene of Tresanti (Florence, Tuscany) in Italy (NHM Inv. A2576); dorsal view, ventral view, apical view. M. Harzhauser & G. C. Kronenberg, 2007 Page 123 Strombus coronatus Defrance, Rutsch, 1936:34—35. {1] Strombus coronatus var. compressonana Sacco, Ertinal-Erent6z, 1958:38, pl. 4, figs. 2-3. [2] Strombus coronatus Defrance, Glibert, 1963:219. [3] Strombus coronatus Defrance, Compagnoni, 1964:259, fig. 6. [4] Strombus coronatus Defrance, Moroni & Paonita, 1964:12. [5] Strombus coronatus Defrance, Hecht et al., 1964:451. [6] Strombus (Strombus) coronatus Defrance, Symeo- nides, 1965:256, pl. 32, fig. 1. [7] Strombus coronatus Defrance, Palla, 1967:959, pl. V2, ville, 7. [8] Strombus coronatus Defrance, Mastrorilli, 1969:pl. Dp 1s, Ze [9] Strombus coronatus Defrance, Beneventi & Piccoli, 1969:17, pl. 2, fig. 1. [10] Strombus (Strombus) coronatus Defrance, Mala- testa, 1974:219, pl. 17, figs. 1-7. [11] Strombus (Strombus) coronatus Defrance, Pavia, 1975:112, pl. 5, figs. 1-4. [12] Strombus coronatus Defrance, Fekih, 1975:111, pl. 33, fig. 1. [13] Strombus coronatus Defrance, Meco, 1977:56, pl. 14, fig. 2, pl. 15, fig. 2, pl. 16, figs. 1—2, ete. [14] Strombus coronatus Defrance, Martinell, 1979:123, pl. 3, figs. 5—6. [15] Strombus coronatus (Defrance), Akbulut, 1980:5. [16] Strombus (Strombus) coronatus Defrance, Bré- bion, 1983:165 (?, see further below). [17] Strombus coronatus var. percoronata Sacco, Fererro-Mortara et al., 1984:138, pl. 21, figs. 2a—2c. [17] Strombus coronatus var. perspinosonana Sacco, Fererro-Mortara et al., 1984:139, pl. 21, figs. 6a—6b. [17] Strombus coronatus var. compressonana Sacco, Fererro-Mortara et al., 1984:139, pl. 21, figs. 7a—7b. [18] Strombus coronatus (Defrance), Moscatelli, 1987:18. [19] Strombus coronatus Defrance, Cavallo & Re- petto, 1992:58, fig. 101. [20] Strombus coronatus (De France), Falconer, 1996:67—-68, figs 1-9, 69 figs 1—9. [21] Strombus coronatus, Gregor et al., 1998:13 middle fig., specimen on right, 13 bottom fig. [22] Strombus coronatus, Ivanov et al., 2001:112 bottom figure, specimen on left. [23] Strombus coronatus (Defrance), Islamoélu, 2002:53, 54. [24] Strombus (Strombus) coronatus Defrance, Lan- dau et al., 2004:63, pl. 14, fig. 6. non Persististrombus coronatus (Defrance, 1827) [but used as presumed valid name as required by ISCN Article 23.9.1.2.] [25] Strombus (Str.) coronatus Defrance, Sieber, 1958:141 (= ex gr. Persististrombus exbonellii Sacco, 1893). [26] Strombus coronatus Defrance, Strausz, 1966:222, figs. 102-103 (= ex gr. Persististrombus lapugyensis- exbonellii Sacco, 1893). [27] Strombus coronatus Defrance, Baldi, 1973:2705, pl. 34, figs. 7-8. (unnamed Persististrombus). [28] Strombus coronatus Defrance, Steininger & Baldi, 1975:345, pl. 3, fig. 6. (unnamed Persististrombus). [29] Strombus (Strombus) coronatus Defrance, Tanar, 1985:22, pl. 1, fig. 4 (= Melongena cornuta Agassiz, 1843). [30] Strombus (Strombus) coronatus Defrance, Niko- lov, 1993:69, pl. 3, figs. 7-8 (= “‘Euprotomus” schroeckingeri Hérnes in Hoernes & Auinger, 1884). [31] Strombus (Strombus) coronatus Defrance, Schultz, 1998:60, pl. 23, fig. 6 (= ex gr. Persistis- trombus lapugyensis Sacco, 1893). [32] Strombus coronatus (Defrance), Arpad, 2003:11 (= ex gr. Persististrombus bonellii Brongniart, 1823). [33] Strombus coronatus (Defrance), Islamoglu & Taner, 2003:22 (= ex gr. Persististrombus bonellii Brongniart, 1823). Note that the suggested affiliations are only prelim- inary; details will be provided in Harzhauser and Kronenberg in prep. Defrance (1827:124) based his description of Strom- bus coronatus on figures in Walch (1768, pl. 38 figs 1—2; attributing the work to Knorr) and added ‘On trouve des coquilles de cette espéce aux environs de Sienne.”’ (One finds shells of this species near Siena). We conclude that Defrance’s description is based on both the illustration by Walch and by specimens from near Siena (Italy) Defrance had seen prior to his description. These specimens are best considered syntypes of Strombus coronatus (ICZN recommendation 73F). Dance (1986:209) gives infor- mation on the Defrance collection, stating it is present in the Musée d’Histoire Naturelle, Caen (France), and some shells in Geneva (Switzerland). According to Cleevely (1983), the Defrance collec- tion in Caen was destroyed in 1944. It is however possible that parts of the Defrance collection, such as parts of the Coelentrata (see Cleevely, 1983), survived this bombing and the whereabouts of possible remains are presently unaccounted for. Nevertheless, Dr. Jean- Philippe Rioult, Université de Caen confirmed to Mr. Franck Frydman (email 13 November 2007) that the Defrance collection indeed was destroyed (‘‘... mais cette collection a bien été détruite en totalité lors du bombardement incendiaire des locaux du Museum d’Histoire Naturelle de Caen le 7 juillet 1944.) and added “A moins d’un miracle (...) il ne faut pas compter retrouver d’échantillons de cette collection.” Dr. Yves Finet, Museum d Histoire Naturelle, Page 124 Geneve (MHNG), informed us that there are no specimens of S. coronatus present in that museum (email 25 Oct. 2007). Rutsch (1936) claimed that the specimen present in the collection of the Naturhistorisches Museum Basel (NMB), coll. nr. 93/1790, 1s the specimen illustrated by Walch. However, comparison of images of the Basel specimen, kindly made available by Mr. Arne Ziems, NMB, here reproduced (Figures 3—5), clearly demon- strates that these are different specimens. Also the accompanying label (Figure 6) makes a provenance from the Walch collection quite improbable. Therefore, the only currently available syntype 1s the specimen illustrated by Walch (1769), and all other syntypes are considered lost. Meco (1977) distinguished S. coronatus from S. latus Gmelin, 1791 (as S. bubonius Lamarck, 1822) based on morphometrics, without fixing a type specimen for S. coronatus. Prior to Meco’s (1977) publication, S. coronatus had been often confused with other species of the Miocene to Recent which we allocate to Persististrombus. Even after Meco’s (1977) paper (see listing above) considerable confusion about the identity of S. coronatus remains. To unequivocally stabilize the identity of S. corona- tus we hereby designate the specimen illustrated in Walch (1768, pl. 38, figs 1-2), here re-illustrated (Figures 1-2), as lectotype of Strombus coronatus Defrance, 1827. Description: Protoconch of all available specimens poorly preserved, with about 3 smooth, moderately convex whorls. Shell of Pliocene specimens very thick and heavy; Tortonian ones are generally slightly less robust. Spire low, concave, with an average apical angle of 70.8° (n = 47, s = 11.0; Table 1) [apical angle = angle of spire whorls; body whorl angle = angle between the flanks of the terminal part of the last whorl; body whorl height = height of last whorl from anterior tip to the suture between last whorl and last spire whorl in apertural view]. Size of adults usually up to 110 mm, but giant representatives may attain sizes up to 155 mm; dwarf forms are also common. About eight prominent triangular shoulder spines on body whorl and evident on the spire whorls as sutural excrescences which produce a stellate (““coronate’’) pattern in apical view. Two more spiral rows of knobs on the body whorl; middle one often reduced or absent while anterior one predominates. Outer lip solid and strongly thickened. Columellar callus thick, covering the base completely in fully grown adults. Often wing extends apically above the suture between the penulti- mate and body whorls. Stromboid notch deeply incised and regularly U-shaped. A typical specimen of S. coronatus is illustrated here (Figures 7-9), Remarks: Herein we treat only Late Miocene to Early The Veliger, Vol. 50, No. 2 Pliocene specimens as Persististrombus coronatus (De- france, 1827). Several morphologically similar speci- mens known from Early to Middle Miocene deposits of the Paratethys Sea usually have been treated as Strombus coronatus Defrance. However, these shells differ in their stromboid notch-morphology (e.g., shallower and less well defined margins) and/or in their higher early spire. Sacco (1893) recognized these differences and introduced several new species names for such specimens. These “‘coronatus-like’’ morphs represent independent, iterative developments within a Persististrombus lineage which comprise a_herein- proposed preliminary concept: the Persististrombus lapugyensis-exbonellii group. A slender counterpart of iterative but unrelated developments is represented by the extant Persististrombus granulatus (Swainson, 1822) and the Middle Miocene Persististrombus exbonellii (Sacco, 1893). Occurrences of Persististrombus in the Pontilevian fauna (Middle Miocene) of the Loire Basin described by Glibert (1949) as Strombus coronatus might also belong to this group. A detailed analysis of the Middle Miocene strombids will be presented elsewhere (Harzhauser & Kronenberg in prep.). Distribution: The earliest record of this species is mentioned by Brébion (1983) from the Middle or Late Miocene of Angola. Unfortunately, Brébion (1983) did not provide a description or an illustration. Therefore, this interesting occurrence has to be treated with caution. If the identification is correct, then the West African Miocene occurrence of Persististrombus cor- onatus suggests that this species is a West African element, which later invaded the Mediterranean Region. There, it does not appear before the Tortonian, where it is known from Italy and Turkey (Sacco, 1893; Stchepinsky, 1939, 1946). It is apparently absent from the Mediterranean during the Messinian but flourished in this bioprovince in the Zanclean and the early Piacenzian. During this warm period the species is recorded from Portugal, Spain, France, Italy, Greece, Turkey, Syria, Libya, Tunisia, Morocco and the Canary Islands (Pereira da Costa, 1866; Almera & Bofill, 1886; Serres, 1829; d’Ancona, 1871; Sacco, 1893; Gignoux, 1913; Eriinal-Erent6z, 1958; Symeonides, 1965; Roman, F. 1940; Fekih, 1975; Lecointre, 1952; Landau et al., 2004; Meco, 1977). Persististrombus coronatus disappears from the Mediterranean Sea completely with the onset of the Late Pliocene cooling (Landau et al., 2004) and seems to be extinct thereafter. CONCLUSIONS Despite its homonymy with a cerithiid described by Roding (1798), Strombus coronatus Defrance, 1827 can be conserved as a name for a Miocene to Pliocene strombid species. A review of the literature fulfills the requirements of the ICZN. Moreover, recent studies of M. Harzhauser & G. C. Kronenberg, 2007 Table | Page 125 Measurements of 47 specimens of Persististromus coronatus (Defrance, 1827) from the collections of the Natural History Museum Vienna (NHM) and Naturalis - Nationaal Natuurhistorisch Museum, Leiden (RGM). =) au OMNADUNAKRWN Ke locality Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Muezzinler - Turkey Bolognese - Italy Bordighera - Italy Bucciana - Italy Toscana - Italy Toscana - Italy Asti - Italy Asti - Italy Asti - Italy Castell Arquato - Italy Castell Arquato - Italy Castell Arquato - Italy Castell Arquato - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Ceriale Rio Torsero - Italy Siena - Italy Siena - Italy Siena - Italy Siena - Italy Tresanti - Italy Tresanti - Italy Tresanti - Italy Tresanti - Italy Tresanti - Italy Sicily Sicily Fuerteventura - Canary Islands Fuerteventura - Canary Islands Fuerteventura - Canary Islands dating Late Miocene Late Miocene Late Miocene Late Miocene Late Miocene Late Miocene Late Miocene Late Miocene Late Miocene Late Miocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene Early Pliocene height (mm) width (mm) aperture height (mm) N I 19 body whorl height (mm) WNN WN m- 080 N apical angle body whorl angle WW Nom) 4B Page 126 Kronenberg & Lee (2007) have shown that this taxon is assignable to the strombid genus Persististrombus, which affiliation unites a conspicuous amphi-Atlantic Neogene species flock. The fossil record suggests that Persististrombus coronatus might have its roots in the Miocene of West Africa. During the Tortonian it managed to invade the Mediterranean Sea. The Messinian crisis forced the species to retreat from the Mediterranean, and it might have found a refuge in the Eastern Atlantic. During the early Pliocene warming it became very abundant throughout the Mediterranean, being recorded from nearly all coasts. Its final extinction was related to the Pliocene cooling. This strict species concept shows that the Early and Middle Miocene populations of Central Europe, erroneously synonymized with P. coronatus in the literature, represent ‘‘coronatus-like’” but unrelated morphs of the Persististrombus lapugyensis-exbonellii group. Acknowledgments. We thank Frank Wesselingh (Naturalis, Leiden) for support during studies in the collection in Leiden and providing some literature. Dr. Birgit Gaitzsch (TU Freiberg, Germany) kindly helped to search for the strombid illustrated by Walch (1768) in the collection in her custody, and Mr. Willem Faber (The Hague, The Netherlands) is acknowledged for his kind support with the literature. We thank Dr. Yves Finet (MHNG) for providing information about the collection in his custody, Mr. Arne Ziems (NMB) for making images of the supposed Walch specimen available, Mr. Franck Frydman (Paris, France) and Dr. Jean-Philippe Rioult (Universté de Caen) for their efforts to gather information on the Defrance collection. GCK wants to thank Ms. Marianne Matthijssen for her abiding support. Dr. Harry G. Lee, Jacksonville, Florida, USA, corrected the English for us. This study contributes to the FWF-Project P-18189-N10: Biogeographic Differentiation and Biotic Gradients in the Western Indo-Pacific during the Late Oligocene to Early Miocene. LITERATURE CITED AKBULUT, A. 1980. 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The Veliger 50(2):129-148 (June 20, 2008) THE VELIGER © CMS, Inc., 2007 Phiocene and Pleistocene Fissurella Bruguiere, 1789 (Gastropoda: Fissurellidae) from Southern Peru THOMAS J. DEVRIES' Burke Museum of Natural History and Culture, University of Washington, Seattle, WA 98195, USA Abstract. Twelve species of fossil Pliocene and Pleistocene keyhole limpets, Fissure/la (Fissurella), from southern Peru are reported, including four new species: Fissurella aranea, sp. nov., F. geoglypha, sp. nov., F. mcleani, sp. nov., and F. persica, sp. nov. These species and others from Peru and Chile are assigned to several morphological species groups that are nearly congruent with clades defined by DNA sequences (Olivares, 2006). Fissurella (Fissurella) probably appeared in western South America during the late Miocene or soon thereafter, possibly from the western Pacific Ocean or South Africa, with major clades in existence in Peru by the late early Pliocene. A late Pliocene extinction of some fissurellid species and the subsequent addition of new and often larger species in Peru, Chile, and Argentina has produced the modern diverse Fissurella (Fissurella) fauna. INTRODUCTION In a systematic treatment of South American keyhole limpets assigned to Fissurella (Fissurella) Bruguiere, 1789, McLean (1984a) lamented their limited fossil record in western South America, a record comprised only of Philippi’s (1887) description of one Pliocene species from Chile, F. concolor Philippi, 1887, Herm’s (1969) list of Pleistocene species and passing mention of Pliocene Fissurella from Chile, and McLean’s own figures of Pliocene F. concolor and Fissurella sp., cf. F. crassa Lamarck, 1822, both also from Chile. This article augments that sparse record with an account of Fissurella in Pliocene and Pleistocene strata from southern Peru, including the extant F. maxima Sowerby, 1835, F. limbata Sowerby, 1835, F. crassa, F. latimarginata Sowerby, 1835, and the extinct F. concolor, which were found in a Pliocene outcrop near the coastal town of Chala, as were two new species, F. persica, sp. nov., and F. aranea, sp. nov. A specimen of F. pulchra Sowerby, 1835, was found in a nearby Pliocene deposit and a worn specimen of F. cumingi Reeve, 1849, was discovered on a nearby middle Pleistocene marine terrace. A Pliocene exposure above the Rio Acari has yielded specimens of F. concolor, F. persica, and the extant F. peruviana Lamarck, 1822. Pliocene sandstones near Yauca produced single specimens of the extinct F. mcleani, sp. nov., and F. geoglypha, sp. nov. The northernmost record for any South American Fissurella (Fissurella) is formally reported, a Pliocene specimen of F. peruviana from northern Peru. ‘Mailing address: Box 13061, Burton, WA 98013 USA GEOLOGY The Cenozoic stratigraphy of southern Peruvian forearc basins (Figure 1) was reviewed by DeVries (1998). Pliocene marine deposits of the Pisco and La Planchada formations include bioclastic sandstones and balanid coquinas (Beaudet et al., 1976; Muizon & DeVries, 1985), the remnants of littoral deposits that lapped onto pre-Eocene crystalline platforms or accumulated at the foot of steep cliffs of the Andes Cordillera. Two outcrops of Fissurella-bearing Pliocene strata warrant mention. Bioclastic debris of the La Planchada Formation crops out along sweeping curves of the Panamerican Highway southeast of Chala where the road descends to the strandline at Playa Huacllaco (Figure 2). The sediments accumulated in high-energy foreshore and rocky intertidal environments in front of a rugged coastline of igneous rock (DeVries, 2003). The age of the Huacllaco beds is constrained by basal beds containing specimens of the muricid gastropods Con- cholepas nodosa MoOricke, 1896, Acanthina triangularis DeVries, 2003, and Herminespina mirabilis (MoOricke, 1896), which collectively indicate a late early to early late Pliocene age (DeVries & Frassinetti, 2003), and by the uppermost and oldest marine terrace, whose elevation and largely extant taxa, including Acanthina unicornis (Bruguiere, 1789) and Concholepas conchole- pas (Bruguicre, 1789), suggest a latest Pliocene age (Muizon & DeVries, 1985; DeVries, 1995, 2000, 2003). Four lithologic units were designated in the Huacllaco section: Unit I (crossbedded balanid coquina, late early Pliocene), Unit II (bioclastic sandstone and polychaete reefs, late early or early late Pliocene), Unit III (ferruginous cobble-bearing bioclastic sandstone, early late Pliocene), and Unit IV (conglomerates and massive coquina beds, late late Pliocene). Page 130 Sacaco ao are SSN Basin ~*~ © 0 100 km Figure 1. Location of forearc basins (Pisco, Sacaco, Ca- mana) along the coast of southern Peru. The second notable outcrop lies east of Chauvina, set back from the southeastern rim of the Rio Acari, where small knobs of bedded strata constitute a condensed section of lower Pliocene to uppermost Pliocene bioclastic sandstone and coquina (Figure 3). The Acari section has a basal lag of igneous boulders with mollusks (Herminespina saskiae DeVries & Vermeij, 1997, Trophon carlosmartini DeVries, 2005, and Xanthochorus ochuroma DeVries, 2005) that indicate an early Pliocene age (DeVries & Vermeij, 1997; DeVries, 2005a, 2005b). Beds five and six meters above the boulders contain Concholepas kieneri Hupé, 1854, and Anadara aff. A. chilensis (Philippi, 1887), as well as the aforementioned muricids, collectively indicating a somewhat later early Pliocene age (Muizon & DeVries, 1985; DeVries, 1995, 2000). Beds nine and 11 m above the boulders with Concholepas camerata DeVries, 2000, Stramonita chocolata (Duclos, 1832), Xanthochorus xuster DeVries, 2005, and Xanthochorus cassidiformis (Blainville, 1832), indicate a late Pliocene age (DeVries, 2000, 2005a, 2007). An overwhelming preponderance of extant molluscan taxa (e.g., Concholepas concholepas and Xanthochorus cassidiformis) in the uppermost coquinas, coinciding with the most elevated and oldest marine terrace, indicate a latest Pliocene age (Muizon & DeVries, 1985). MATERIALS AND METHODS Specimens of fossil Fissurella described in this study were found by the author. Most fossil examples had lost their aragonitic inner layer, so features of the external calcitic layer (shape, radial and concentric sculpture, ray patterns, margin width and coloration) The Veliger, Vol. 50, No. 2 became the principal means for diagnosing species, with greatest emphasis placed on radial sculpture. Comparative fissurellid material came from the Natural History Museum of Los Angeles County (LACM) and the author’s personal collection. Selected citations are given for known species, emphasizing those that post-date McLean (1984a). Locality and sample descriptions are listed in the appendix. Lengths (L), widths (W) and heights (H) are measured in millimeters. Dimensions of broken spec- imens are enclosed by parentheses. Some figured specimens are coated with ammonium chloride; others are shown with transmitted light to reveal color patterns. Types and other numbered specimens are deposited at the University of Washington’s Burke Museum of Natural History and Culture in Seattle, Washington (UWBM), the Departamento de Paleon- tologia de Vertebrados, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, in Lima, Peru (MUSM INV) and in the case of one specimen of Fissurella peruviana, the Orton Geological Museum, The Ohio State University (OSU). SYSTEMATIC PALEONTOLOGY Family Fissurellidae Fleming, 1822 Subfamily Fissurellinae Fleming, 1822 Genus Fissurella Bruguiere, 1789 Subgenus Fissure/la Bruguiere, 1789 Type species (by monotypy) Patella nimbosa Linnaeus, 1758. Recent, Caribbean. Discussion: McLean (1984a) included within Fissurella (Fissurella) all fissurellid gastropods with an aragonitic inner shell layer and calcitic outer shell layer, 1.e., 13 South American taxa and two species from the northern hemisphere, F. (Fissurella) volcano Reeve, 1849 (California), and F. (Fissurella) nimbosa (Lin- naeus, 1758) (Caribbean Sea, Brazil). Stuber (1991) supported the monophyly of Fissurella (Fissurella) sensu. McLean (1984a), but Olivares (2006) did not, concluding instead that molecular data showed F. nimbosa to be more closely related to the eastern Atlantic F. (Cremides) schrammi Fischer, 1857, than South American species of Fissurella (Fissurella). Until molecular data become available for F. volcano, McLean’s (1984a) definition of Fissurella (Fissurella) will be utilized here, with the caveat that removing both F. volcano and F. nimbosa from the subgenus would result in a South American subset of Fissurella equivalent to Pérez-Farfante’s (1943) imperfectly diag- nosed Fissurella (Balboaina). T. J. DeVries, 2007 Page 131 IV DV 1032-1 (F latimarginata, F. maxima) DV 1628-6 (F) maxima) UNIT Hl | —DV 1254-Bal 10 (F maxima, limbata) 2—DV 1254-Bal 8, -5 (F maxima, limbata) ee —DV 1254-6, 14 (F concolor, limbata) fe a concolor, CraSSa | UNIT U 2048 Dien UNIT I 10 0 meters [| Cobbles Massive sandstone b * Coquina °.7.-) Pebbly sandstone Bioclastic rippled sandstone ~ <7) Igneous basement Polychaete colony Figure 2. The Huacllaco lithologic section southeast of Chala, type locality of Fissurella aranea, sp. nov. The location map is shown in inset. “1 Bioclastic crossbedded sandstone Page 132 The Veliger Vol; 50; Now? UPPERMOST Sandy coquina of Glycymeris ovata and Discinisca MARINE (brachiopod) with Xanthochorus xuster, Concholepas TERRACE concholepas Sandy coquina of Glycymeris ovata and Discinisca (brachiopod) with Xanthochorus xuster, Concholepas concholepas Bioclastic coarse-grained sandstone with Glycymeris ovata, Discinisca, venerids, Choromytilus, Xanthochorus xuster, Concholepas camerata LATE PLIOCENE DV 923-1E Balanid coquina with Concholepas camerata, DV 923-1 Xanthochorus xuster, Chlamys sp.; base with EB peruviana rounded and angular basement boulders F. persica Balanid coquina with Concholepas kieneri, 1) Xanthochorus ochuroma, Anadara cf. A. chilensis Balanid coquina with Concholepas kieneri, Z. Herminespina saskiae, Anadara cf. A. chilensis = Balanid coquina with Concholepas kieneri, © Herminespina saskiae, Xanthochorus ochuroma, @ Anadara cf. A. chilensis, Chlamys vidali, Piscoacritia al collapsa — Shelly gravel and angular boulders with _ Herminespina saskiae, Xanthochorus ochuroma, > Trophon carlosmartini, Concholepas nodosa < DV 923-1A aa F; persica F. concolor cs to. () \ a COVERED meters = | Boulders and cobbles Polychaete colony SAE Massive sandstone 2 5| Coquina ep Ophiomorpha burrows Figure 3. Lithologic section at the Acari site southeast of the Rio Acari, type locality (DV 923) for Fissurella persica, sp. nov. The location map is shown in inset. Associated mollusks listed in section are discussed in DeVries (2000, 2003, 2005a, 2005b), DeVries & Hess (2004), and DeVries & Vermeij (1997). T. J. DeVries, 2007 Fissurella (Fissurella) maxima Sowerby, 1835 Figures 4-10, 17, 18 Fissurella maxima Sowerby, 1835a, p. 123. Fissurella maxima Sowerby, 1835b, p. 3, fig. 18. Fissurella maxima Sowerby, 1835. McLean, 1984a, p. 25, figs. 51-63. Fissurella maxima Sowerby, 1835. Oliva & Castilla, 1992, p. 92, fig. 4. Fissurella maxima Sowerby. Alamo & Valdivieso, 1997, p. 7, fig. 12. Fissurella maxima Sowerby, 1835. Guzman et al., 1998, p. 29, with figure. Fissurella maxima Sowerby, 1834 [sic]. Forcelli, 2000, p. 48, fig. 23. Diagnosis: Shell large. Ribs strong, wide, well differen- tiated by size, corrugated to imbricate. Purplish rays broad. Margin coarsely crenulated. Rays penetrate entire calcitic layer. Discussion: Small specimens with characters of Fissur- ella maxima, including coloration, were found in bioclastic sandstones in the upper part of the Huacllaco section (Figure 2) and assigned a late Pliocene age based on the presence of associated molluscan taxa [Prisogaster valenciai DeVries, 2006, Acanthina unicor- nis, Chorus transitional between C. giganteus (Lesson, 1830) and C. grandis (Philippi, 1887), Concholepas camerata, Xanthochorus xuster (DeVries 1997, 2000, 2003, 2005a, 2006)). Two Peruvian specimens of Fissurella maxima exhibit oval external scars from the epibiotic limpet, Scurria variabilis (Sowerby, 1839) [= Scurria parasitica (Orbigny, 1841); see Espoz et al., 2004] (Figure 8). Such scars are most often found on Recent specimens of F. crassa and F. limbata (Figures 22, 35). Material: MUSM INV 164, DV 401-1, Recent, L 44.4, W 30.9, H 10.1; MUSM INV 165, DV 1032- 1, late Pliocene, L 37.6, W 27.5, H 7.7; UWBM 98437, Lomas, Recent, L 87.7, W 59.4, H 22.0; UWBM 98438, Lomas, L 75.9, W 48.7, H 17.5; UWBM 98439, Lomas, L 55.3, W 36.9, H 13.2; UWBM 98440, Lomas, L 51.6, W 31.7, H 10.6; UWBM 98441, DV 1372-1, Recent, L 60.3, W 35.8, H 13.7; UWBM 98442, DV 1372-1, L 65.7, W 42.8, H 14.7; UWBM 98443, DV 1628-6, late Pliocene, L 43.5, W 29.3, H 10.6; UWBM 98444, DV 1254-Ball0, late Pliocene, L 16.3, W 10.4, H 4.5; UWBM 98482, DV 1418-1, late Pliocene, L 32.6, W 21.5, H 5.3; UWBM 98483, DV 1418-1, L (12.6). Occurrence: Late Pliocene: southern Peru. Pleistocene: central Chile. Recent: north-central Peru to central Chile. Page33 Fissurella (Fissurella) concolor Philippi, 1887 Figures 11—16, 19 Fissurella concolor Philippi, 1887, p. 98, pl. 58, fig. 8. Fissurella concolor Philippi, 1887. McLean, 1984a, Oo Wo 1s IF Diagnosis: Shell medium-sized, elongate, tapered. Ribs strong, well differentiated by size, weakly corrugate or smooth. Margin moderately wide, coarsely crenulated. Description: Shell length up to 60mm, elongate, strongly tapered anteriorly. Height low. Sides of shell elevated. Ribs prominent, sharply rounded, smooth to slightly imbricate, differentiated into primary ribs, some bifurcated, usually with three intervening second- ary ribs, the medial being stronger; smaller specimens with one intervening secondary rib between primary ribs. Margin moderately wide, crenulated by ribs. Rays narrow, purplish, generally coinciding with primary ribs; anterior rays weak or absent. Aragonitic inner layer missing. Foramen anterior to center, shape unknown. Discussion: Fissurella concolor was first described from Pliocene beds near Mejillones, Chile by Philippi (1887) and figured again by McLean (1984a). Peruvian specimens, like those from Chile, are elongate and have ribs that are more sharply raised and _ less imbricate than those on specimens of F. maxima. The oldest example from Peru came from the lowest boulder bed of the Acari locality (DV 923-1la) together with F. persica, sp. nov., and mollusks indicating an early Pliocene age [Acanthina triangularis DeVries, 2003, Concholepas nodosa, C. kieneri, and Hermine- spina saskiae (DeVries, 1995, 2000, 2003; DeVries & Vermeij, 1997)]. Other specimens were found in the lowest cobble-rich bioclastic sandstone in the Huacl- laco section (DV_ 1254-14); associated mollusks (Acanthina transitional between A. triangularis and A. unicornis; Concholepas camerata; Xanthochorus xuster) indicate an early late Pliocene age. Higher in the Huacllaco section the only strongly ribbed Fissurella specimens are referable to F. maxima. Material: MUSM INV 171, DV _ 1032-2, early late Pliocene, L 35.9, W 22.1, H 7.0; MUSM INV 172, Huacllaco, early late Pliocene, L (26.2), W 16.9, H 5.9; MUSM INV 173, DV 1254-14, early late Pliocene, L 28.0, W 16.4, H 3.3; UWBM 98463, DV 923-la, early Pliocene, L (11.1), W 9.1, H 2.0; UWBM 98464, Huacllaco, late Pliocene, L 47.9, W 29.1, H 10.8; UWBM 98465, DV 1032-2, L 24.5, W 14.5, H_ 5.2; UWBM 98466, DV 1032-2, L 19.6, W 16.9, H (5.1); UWBM 98467, DV 1254-6, late Pliocene, L 19.3, W The Veliger, Vol. 50, No. 2 Figures 4-10, 17, 18. Fissurella (Fissurella) maxima Sowerby, 1835. Figure 4. Figure 5. Figure 6. Figure 7. UWBM 98443, DV 1628-6. Late Pliocene. Transmitted light showing rays. Length is 43.5 mm. UWBM 98443. Dorsal view. UWBM 98443. Ventral view, aragonitic inner layer missing. UWBM 98441, DV 1372-1. Recent. Transmitted light showing rays. Length is 60.3 mm. Te Jo IDEWmES, AVOY Page 135 11.5, H 4.1; UWBM 98468, DV 1254-14, L 29.6, W 18.4, H 3.9; UWBM 98469, DV 1254-14, L (37.9). Occurrence: Early to early late Pliocene: southern Peru. Pliocene: north-central Chile. Fissurella (Fissurella) aranea, sp. nov. Figures 20, 21, 34 Diagnosis: Shell small, elongate-quadrate. Height low. Ribs sharply defined, alternating primary and second- ary. Margin wide. Description: Shell small, less than 30 mm_ long, elongate-quadrate, slightly tapered anteriorly. Height low. Ends slightly raised. Sculpture of well defined, closely spaced, narrow ribs, alternatingly primary and secondary; moderately imbricate at intersections with strong growth lines. No coloring preserved. Margin wide, irregularly scalloped. Foramen probably slightly anterior to center. Discussion: Specimens of Fissurella aranea differ from contemporaneous small specimens of F. persica, sp. nov., by being more quadrate and having ribs that alternate regularly in size, in contrast with fine equally sized ribs on specimens of F. persica. Ribs on specimens of F. concolor and F. maxima are coarser and alternate less regularly. Etymology: Latin noun ‘aranea, meaning ‘spider net,’ referring to the intersecting ribs and concentric growth lines on this species. Type Locality: DV 1254, section along Panamerican Highway above Playa Huacllaco, 27 m in measured section (see Appendix). Material: UWBM 98479, DV 1254-Bal5, holotype, late early Pliocene; L 27.4, W 16.7, H (4.9); UWBM 98480, DV 923-le, early late Pliocene, L (17.9), W 13.2, H (2.7). — Occurrence: Late early Pliocene to early late Pliocene: southern Peru. Fissurella (Fissurella) limbata Sowerby, 1835 Figures 22—33 Fissurella limbata Sowerby, 1835a, p. 123. Fissurella limbata Sowerby, 1835b, p. 3, figs. 42, 66, 74. Fissurella limbata Sowerby, 1835. McLean, 1984a, p. 55, figs. 212-224. Fissurella limbata Sowerby, 1835. Oliva & Castilla, 1992, p. 7, fig. 10. Fissurella limbata Sowerby. Alamo & Valdivieso, 1997, p. 93, fig. 10. Fissurella limbata Sowerby, 1835. Guzman et al., 1998, p. 28, with figure. Fissurella limbata Sowerby, 1835. Forcelli, 2000, p. 49, fig. 25. Diagnosis: Shell medium-sized. Ribs broad, subdued, some with weak secondary ribs; ribs often obsolete towards foramen. Margin wide. Calcitic outer layer with translucent veneer. Discussion: Modern specimens of Fissurella limbata have ribs that are less well defined than those of F. maxima. Such specimens were encountered in bioclastic sandstones in the upper half of the Huacllaco section near Chala (DV 1254-14), together with specimens of F. concolor. One specimen (Figure 27) has a nearly full suite of primary, secondary, and tertiary ribs extending to the foramen, such as is seen on specimens of F. maxima, although the ribs are subdued and lack the imbricate texture of the latter species. Rays are seen to penetrate the entire calcitic outer layer, but perhaps only because the purple color of the flattened margin is faded. Associated mollusks indicate a late Pliocene age. Not one Pliocene specimen exhibited the external scar of the epibiotic limpet, Scurria variabilis, which is commonly seen on modern specimens of F. /imbata (Figures 22, 29). Figure 8. MUSM INV 165, DV 1032-1. Late Pliocene. Dorsal view; arrow marks scar from Scurria limpet. Length is 37.6 mm. Figure 9. MUSM INV 165, ventral view. Figure 10. UWBM 98440, Lomas. Recent. Dorsal view. Length is 51.6 mm. Figure 17. UWBM 98443, lateral view, anterior to right. Figure 18. UWBM 98441, lateral view, anterior to right. Figures 11-16, 19. Fissurella (Fissurella) concolor Philippi, 1887. UWBM 98464, Huacllaco. Early late Pliocene. Dorsal view. Length is 47.9 mm. _ Figure 11. Figure 12. MUSM INV 171, DV 1032-2. Late early Pliocene. Dorsal view. Length is 35.9 mm. Figure 13. MUSM INV 171, transmitted light. Figure 14. UWBM 98465, DV 1032-2. Early late Pliocene. Dorsal view. Length is 24.5 mm. Figure 15. MUSM INV 173, DV 1254-14. Early late Pliocene. Dorsal view. Length is 28.0 mm. Figure 16. UWBM 98464, ventral view. Figure 19. UWBM 98464, lateral view, anterior to right. Page 136 The Veliger, Vol. 50, No. 2 Figures 20, 21, 34. Fissurella (Fissurella) aranea, sp. nov. UWBM 98479, DV 1254-Bal5. Holotype. Late early Pliocene. Length is 27.4 mm. Figure 20. Dorsal view. Figure 21. Ventral view. Figure 34. Lateral view, anterior to right. Figures 22-33. Fissurella (Fissurella) limbata Sowerby, 1835. Figure 22. MUSM INV 167, DV 810-1. Holocene. Dorsal view. Length is 60.8 mm. Arrow marks scar from Scurria limpet. Figure 23. MUSM INV 167, ventral view. Figure 24. MUSM INV 170, Hucallaco. Early late Pliocene. Dorsal view. Length is 37.6 mm. Figure 25. MUSM INV 170, ventral view. T. J. DeVries, 2007 Material: MUSM INV 163, DV 1372-1, Recent, L 63.4, W 43.7, H 15.3; MUSM INV 167, DV _ 810-1, Holocene, L 60.8, W 43.6, H 18.7; MUSM INV 168, DV 1254-5, late Pliocene, L 62.8, W 42.1, H 17.0; MUSM INV 169, DV 1254-14, early late Pliocene, L (28.5); MUSM INV 170, lowest Huacllaco cobbles, 37 m, early late Pliocene, L 37.6, W 24.9, H 6.5; UWBM 98452, Lomas, Recent, L 50.4, W 33.7, H 12.9; UWBM 98453, Lomas, L 48.8, W 33.6, H 11.9; UWBM 98454, DV 810-1, L 72.2, W 49.8, H 18.8; UWBM 98455, DV 1254-6, late Pliocene, L 53.9, W 37.5, H (15); UWBM 98456, DV 1254-14, L 42.6, W 30.1, H 11.3; UWBM 98460, DV_ 1254-10, late Pliocene, L 36.4, W 24.4, H 8.2; UWBM 98461, DV 1254-14, L 34.6, W 21.9, H 6.2; UWBM 98462, DV 1254-14, L 36.6, W 23.9, H 8.7; UWBM 98457, DV 1372-1, L 36.9, W 23.9, H 9.0; UWBM 98458, DV 1372-1, L 38.6, W 25.9, H 8.9; UWBM 98459, DV 1372-1, L (22.9), W 16.8, H 7.0; UWBM 98478, DV 1254-2, early late Pliocene, L (27.7). Occurrence: Late Pliocene: southern Peru. Recent: north-central Peru to Chiloé, Chile. Fissurella (Fissurella) crassa Lamarck, 1822 Figures 35—37 Fissurella crassa Lamarck, 1822, 6(2), p. 11. Fissurella crassa Lamarck, 1822. Sowerby, 1835b, p. 1, figs. 9, 11. Fissurella crassa Lamarck, 1822. McLean, 1984a, p. 58, figs. 225-237. Fissurella crassa Lamarck, 1822. Oliva & Castilla, 1992, p- 92, fig. 5. Fissurella crassa Lamarck. Alamo & Valdivieso, 1997, p. 6, fig. 11. Fissurella crassa Lamarck, 1822. Guzman et al., 1998, p. 27, with figure. Fissurella crassa Lamarck, 1822. Forcelli, 200, p. 46, fig. 13. Diagnosis: Shell medium-sized to large, elongate, with upturned ends and margins; margin with fimbriate edge. Ribs and rays poorly developed. Foramen elongate. Co Page 137 Discussion: Specimens of Fissurella crassa are more elongate and less tapered than those of F. /imbata and have nearly obsolete ribs. A fragment of Fissure/la from the lower half of the Huacllaco section (Figure 37) has the same uniquely scalloped, fimbriate, upturned margin as specimens of extant F. crassa and so is referred to this species. Material: MUSM INV 166, DV 810-1, Holocene, L 61.1, W 36.3, H 13.3; UWBM 98447, DV 1372-1, Recent, L 63.6, W 35.9, H 14.7; UWBM 98448, Lomas, Recent, L 52.3, W 31.1, H 11.3; UWBM 98449, Lomas, L 42.6, W 24.1, H 9.6; UWBM 98450, DV 810-1, L 62.8, W 39.6, H 15.4; UWBM 98451, Huacllaco, 39 m, early late Pliocene, L (19.7). Occurrence: Early late Pliocene: southern Peru. Late Pliocene: Chile. Recent: north-central Peru to Chiloé, Chile. Fissurella (Fissurella) peruviana Lamarck, 1822 Figures 38—42 Fissurella peruviana Lamarck, 1822, p. 15. Fissurella peruviana Lamarck, 1822. McLean, 1984a, p. 21, figs. 31—SO. Fissurella peruviana Lamarck, 1822. Alamo & Valdi- vieso, 1997, p. 7, fig. 13. Fissurella peruviana Lamarck, 1822. Guzman et al., 1998, p. 29, with figure. Fissurella peruviana Lamarck, 1822. Forcelli, 2000, p. 48, fig. 22. Diagnosis: Shell small, profile high. Fine primary and secondary ribs. Color usually charcoal gray; rays obscured. Margin narrow. Foramen small, oval. Discussion: Specimens with the high conic shell and fine ribs of Fissurella peruviana were found in a coquina bed nine meters above the valley floor of the Acari outcrop (Figure 3). Associated mollusks (Concholepas camerata, Xanthochorus xuster, Stramonita chocolata) indicate a late Pliocene age (DeVries, 2000, 2005a, 2007). A specimen (OSU 38157) assigned herein to Fissur- ella. peruviana was found in the gravel-rich cross-bedded sandstones of the basal Taime Formation in northern Peru (DV 239-11; DeVries, 1986, 1988). Figure 26. UWBM 98456, DV 1254-14. Early late Pliocene. Dorsal view. Length is 42.6 mm. Figure 27. UWBM 98462, DV 1254-14. Dorsal view. Length is 36.6 mm. Figure 28. UWBM 98461, DV 1254-14. Dorsal view. Length is 34.6 mm. Figure 29. UWBM 98457, DV 1372-1. Recent. Dorsal view. Length is 36.9 mm. Arrow marks scar from Scurria limpet. Figure 30. UWBM 98460, DV 1254-10. Late Pliocene. Dorsal view. Length is 36.4 mm. Figure 31. UWBM 98460, transmitted light showing rays. Figure 32. UWBM 98461, lateral view, anterior is to right. Figure 33. UWBM 98462, lateral view, anterior is to left. Page 138 The Veliger, Vol. 50, No. 2 51 Figures 35-37. Fissurella (Fissurella) crassa Lamarck, 1822. Figure 35. MUSM INV 166, DV 810-1. Holocene. Dorsal view. Length is 61.1 mm. Arrow marks scar from Scurria limpet. Figure 36. MUSM INV 166, ventral view. Figure 37. UWBM 98451, Huacllaco. Early late Pliocene. Dorsal view of fragment. Length is 19.7 mm. Figures 38-42. Fissurella (Fissurella) peruviana Lamarck, 1822. T. J. DeVries, 2007 Page 139 The elongate foramen differs from the oval foramen seen on southern Peruvian and Chilean specimens (Figure 38) but the size and arrangement of radial ribs are identical, as is the high conic profile. Material: MUSM INV 161, DV 923-le, early late Pliocene, L (25.3); MUSM INV 162, DV 923-1, early late Pliocene, L (26.0), W 21.5, H (7.4); OSU 38157, DV 239-11, late Pliocene, L 21.4, W 20.7, H 10.8; UWBM 98424, DV 1141-1, Recent, L 40.3, W 34.2, H 16.5; UWBM 98425, DV 1141-1, L 33.4, W 23.9, H 15.4; UWBM 98426, La Mina, Recent, L 33.4, W 24.9, H 12.6; UWBM 98427, DV 599-2, middle Pleistocene, L 31.0, W 22.5, H 10.2; UWBM 98428, DV 1141-1, L 19.9, W 13.8, H 9.3; UWBM 98429, DV 401-1, Recent, lot of 2; UWBM 98430, DV 923-le, early late Pliocene, L (20.6), W 25.3, H 10.4. Occurrence: Early late Pliocene: northern to southern Peru. Recent: north-central Peru to south-central Chile. Fissurella (Fissurella) latimarginata Sowerby, 1835 Figures 43, 44, 47, 48 Fissurella latimarginata Sowerby, 1835a, p. 126. Fissurella latimarginata Sowerby, 1835b, p. 3, fig. 69. Fissurella latimarginata Sowerby, 1835. McLean, 1984a, p. 28, figs. 64-79. Fissurella latimarginata Sowerby, 1835. Oliva & Cas- tilla, 1992, p. 92, fig. 6. Fissurella latimarginata Sowerby. Alamo & Valdivieso, 1997, p. 6. Fissurella latimarginata Sowerby, 1835. Guzman et al., 1998, p. 28, with figure. Fissurella latimarginata Sowerby, 1835. Forcelli, 1835, p. 46, fig. 12. Diagnosis: Shell large, height low to moderate. Color charcoal gray. Ribs fine, poorly differentiated; rays usually obscured. Margin wide. Foramen elongate. Discussion: A specimen of Fissurella latimarginata (Figure 43) was found in the upper part of Unit III of the Huacllaco section (Figure 2), together with the transitional Chorus giganteus/C. grandis and Conchole- pas camerata, both indicative of a late Pliocene age. Another specimen of F. /atimarginata was found at Quebrada Champeque (DV 1251-1), 10m _ above basement rocks and 10 m below the uppermost marine terrace, together with Concholepas camerata, Xantho- chorus xuster, and Acanthina unicornis, which collec- tively indicate a late Pliocene age (DeVries, 2000, 2003, 2005a). Material: MUSM INV 163, DV 1251-1, late Pliocene, L (27.9), 19.2, H 7.8; UWBM 98433, DV 1032-1, late Pliocene, L 49.5, W 35.6, H 9.1; UWBM 98434, DV 1372-1, Recent, L 55.6, W 37.2, H 13.2; UWBM 98435, Lomas, Recent, L 69.5, W 50.2, H 16.4; UWBM 98436, Lomas, 53.5, W 37.5, H 11.8. Occurrence: Late Pliocene: southern Peru. Recent: north-central Peru to central Chile. Fissurella cumingi Reeve, 1849 Fissurella cumingi Reeve, 1849, pl. 3, fig. 17. Fissurella cumingi Reeve, 1849. Hupé, 1854, p. 238. Fissurella cumingi Reeve, 1849. McLean, 1984a, p. 31, figs. 80-94. Fissurella cumingi Reeve, 1849. Oliva & Castilla, 1992, p. 92, fig. 7. Fissurella cumingi Reeve, 1849. Guzman et al., 1998, p. 27, with figure. —_— Figure 38. OSU 38157, DV 239-11. Late Pliocene. Dorsal view. Length is 21.4 mm. Figure 39. Figure 40. UWBM 98424, lateral view, anterior is to left. UWBM 98424, DV 1141-1. Recent. Dorsal view. Length is 40.3 mm. Figure 41. UWBM 98430, DV 923-le. Early late Pliocene. Dorsal view of fragment. Length is 20.6 mm. Figure 42. MUSM INV 162, DV 923-1. Early late Pliocene. Dorsal view of fragment. Length is 26.0 mm. Figures 43, 44, 47, 48. Fissurella (Fissurella) latimarginata Sowerby, 1835. Figure 43. UWBM 98433, DV 1032-1. Late Pliocene. Dorsal view. Length is 49.5 mm. Figure 44. UWBM 98433, ventral view. Figure 47. MUSM INV 163, DV 1251-1. Late Pliocene. Dorsal view. Length is 27.9 mm. Figure 48. MUSM INV 163, lateral view, anterior is to right. Figures 45, 46. Fissurella (Fissurella) persica sp. nov. Figure 45. UWBM 98470, DV 923-la, holotype. Early Pliocene. Dorsal view. Length is 31.9 mm. Figure 46. UWBM 98477, DV 1254-Bal5. Late early Pliocene. Dorsal view. Length is 20.5 mm. Figures 49-54. Fissurella (Fissurella) pulchra Sowerby, 1835. Figure 49. UWBM 98432, DV 1251-1. Late Pliocene. Dorsal view. Length is 52.6 mm. Figure 50. LACM 75-31 a, Chile. Recent. Close-up showing unique mottling. Figure 51. UWBM 98432, close-up showing unique mottling. Figure 52. LACM 75-31 b, Chile. Recent. Transmitted light showing acicular rays. Figure 53. UWBM 98432, ventral view. Figure 54. UWBM 98432, transmitted light showing acicular rays. Page 140 Fissurella cumingi Reeve, 1849. Forcelli, 2000, p. 48, fig. 24. Diagnosis: Shell large, ight colored, with fine ribs and well-defined rays. Discussion: A large broken and worn specimen of Fissurella cumingi was found on a middle Pleistocene terrace (elevation 160 m above sea level) north of Chala. A large modern specimen from Lomas is very similar to the Pleistocene specimen. Material LACM 75-32, Chile, Recent, lot of 31; UWBM 98445, DV 465-1, middle Pleistocene, L (59.3), W 55.0, H 20.3; UWBM 98446, Lomas, Recent, L 77.3, W 55.7, H 25.3. Occurrence: Early middle Pleistocene: southern Peru. Recent: southern Peru to south-central Chile. Fissurella (Fissurella) persica, sp. nov. Figures 45, 46 Diagnosis: Shell small, height low. Ribs fine, weak, generally undifferentiated. Rays broad, forming a suffusion of purple-peach color in transmitted light. Description: Shell small, length less than 40 mm. Oval, sharply tapered anteriorly. Height low. Calcitic outer layer thin. Sculpture of numerous thin ribs; ribs equally sized, barely raised, differentiated in anterior quadrant only. Ground color cream; broad rays of purplish-peach color. Margin flat, narrow, coloring of rays passes through entire thickness of outer layer. Foramen not preserved, probably just anterior to center. Discussion: Specimens of Fissurella persica closely resemble those of the opaquely gray F. /atimarginata with their fine ribs and sharply tapered anterior. They are lighter colored, however, like specimens of F, cumingi, with visible rays and a narrow margin. Their ribs are more uniformly sized than those of both F. latimarginata and F. cumingi. The holotype of Fissurella persica was found in bioclastic gravel near the base of the Acari outcrop (Figure 3). Its occurrence with Trophon carlosmartini, Herminespina saskiae, and Xanthochorus ochuroma indicates an early Pliocene age. Fragments of F. persica were also found in the upper bioclastic sandstones of the Acari locality (early late Pliocene) and Unit I of the Huacllaco outcrop (late early Pliocene). Etymology: Latin noun ‘persica,’ meaning ‘peach,’ for the purple-peach color of the species’ rays under transmitted light. The Veliger, Vol. 50, No. 2 Type Locality: DV 923, knobs southeast of the Rio Acari, east of Chauvina (Figure 3; see Appendix). Material: MUSM INV 174, DV 923-1, early late Pliocene, L (12.9); MUSM INV 175, DV 923-1, L (17.5); UWBM 98431, DV 923-1, early late Pliocene, L (21.8); UWBM 98470, DV 923-la, early Pliocene, holotype, L (31.9), W 25.0, H 7.2; UWBM 98471, DV 1254-Bal5, late early Pliocene, L (25.4); UWBM 98472, DV 923-le, early late Pliocene, L (27.3); UWBM 98473, DV 923-le, L (29.5); UWBM 98476, DV 1254-2, early late Pliocene, L 22.7, W 15.4, H 2.8; UWBM 98477, DV 1254-Bal5, (20.5), W 14.4, H 3.4. Occurrence: Early to early late Pliocene: southern Peru. Fissurella (Fissurella) pulchra Sowerby, 1835 Figures 49-54 Fissurella pulchra Sowerby, 1835a, p. 124. Fissurella pulchra Sowerby, 1835b, p. 3, fig. 24. Fissurella pulchra Sowerby, 1835. McLean, 1984a, p. 63, figs. 254-267. Fissurella pulchra Sowerby, 1835. Oliva & Castilla, 1992, p. 93, fig. 11. Fissurella pulchra Sowerby, 1835. Guzman et al., 1998, p. 30, with figure. Fissurella pulchra Sowerby, 1835. Forcelli, 2000, p. 49, fig. 27. Diagnosis: Shell medium-sized, smooth. Height very low. Surface with purple-and-white streaky mottling. Broad rays superimposed on irregularly spaced acicular rays. Discussion: A single specimen of Fissurella pulchra, distinguished by its exterior mottling and smooth surface, was discovered in late Pliocene deposits north of Chala. Both modern and fossil examples have weakly developed ribs anteriorly. Material: LACM 75-31, Recent, Chile, lot of 10; UWBM 98432, DV 1251-1, L 52.6, W 36.0, H 8.4. Occurrence: Late Pliocene: southern Peru. Recent: north-central Peru to central Chile. Fissurella (Fissurella) mcleani, sp. nov. Figures 55—57 Diagnosis: Shell medium-sized, elongate. Exterior black, smooth. Foramen exceptionally elongate. Ends elevated. Description: Shell medium-sized, length almost 60 mm, elongate, tapered slightly anteriorly; ends of shell elevated. Height low. Surface black, smooth, with rare radial wrinkles marking edges of obsolete radial ribs. Numerous thin reddish rays, only visible, barely, T. J. DeVries, 2007 Page 141 Figures 55-57. Fissurella (Fissurella) mcleani, sp. nov. UWBM 98475, DV 1635-2. Holotype. Early Pliocene. Length is 58.4 mm. Figure 55. Dorsal view. Figure 56. Ventral view showing partial layer of aragonite and very elongate foramen. Figure 57. Lateral view, anterior to right. Figures 58-60. Fissurella (Fissurella) geoglypha, sp. nov. UWBM 98474, Yauca. Holotype. Early Pliocene. Length is 35.0 mm. Figure 58. Lateral view, anterior to right. Figure 59. Dorsal view. Figure 60. Ventral view, showing partial layer of aragonite. Figure 61. Diodora sp. UWBM 98481, DV 923-la. Early Pliocene. Dorsal view of fragment; note squared off posterior to left. Length is 18.4 mm. apically. Margin moderately wide, smooth, with broad obsolete rays producing alternating cream and black- colored intervals. Aragonite layer partially preserved. Foramen situated slightly posteriorly; exceptionally elongate, without constrictions. Discussion: Fissurella (Fissurella) mcleani has an exceptionally elongate foramen and a smooth black surface. Specimens of Fissurella pulchra are equally smooth, but possess a normally elongate foramen and lack the deep black color of F. mcleani. Associated mollusks from the same beds (Anadara cf. A. chilensis, Acanthina obesa DeVries, 2003, Conchole- pas kieneri, Herminespina saskiae, Xanthochorus ochur- oma) at locality DV 1635-1 indicate an early Pliocene age; the presence of X. eripepomus DeVries, 2005, further suggests a late early Pliocene age. Etymology: Named in recognition of James H. McLean (Natural History Museum of Los Angeles County) and his studies on Fissurella (Fissurella). Type Locality: DV 1635, Yauca depression, west of Panamerican Highway, northwest of Yauca (Figure 62; see Appendix). Material: UWBM 98475, DV 1635-2, holotype, late early Pliocene, L 58.4, W 32.6, H 13.3. Occurrence: Late early Pliocene: southern Peru. Fissurella (Fissurella) geoglypha, sp. nov. Figures 58—60 Diagnosis: Shell medium-sized, smooth. Exterior black; cream-colored rays sharply defined, broad. Description: Shell with estimated length of 60 mm, anterior strongly tapered, posterior unknown. Height low. Anterior end of shell elevated. Exterior smooth. Rays sharply defined, broad, penetrate calcitic outer layer. Margin rounded, moderately wide. Aragonitic shell layer partially preserved. Discussion: The holotype of Fissurella geoglypha is more bowed than the smooth flat specimens of F. pulchra and lacks the latter’s characteristic mottling; it is also less quadrate than the holotype of F. mcleani, which lacks any broad rays. Associated mollusks from the Yauca roadcut (Anadara cf. A. chilensis, Concho- Page 142 lepas nodosa, Xanthochorus ochuroma) indicate an early Pliocene age (DeVries, 2000, 2005a). Etymology: Named for radiating geoglyphs that are part of the pre-Columbian Nazca lines in southern Peru. Type Locality: Yauca, roadcut in Panamerican High- way on north side of Rio Yauca (Figure 62; see Appendix). Material: UWBM 98475, Yauca, holotype, late early Pliocene, L (35.0), W (36.5), H 12.1. Occurrence: Late early Pliocene: southern Peru. DISCUSSION There is no consensus regarding the grouping of related species within Fissurella (Fissurella). Pilsbry (1890) recognized four species groups based on shell mor- phology, whereas McLean (1984a) defined three groups according to the relative thickness of the aragonitic inner and calcitic outer layers. Stuber (1991) utilized 34 anatomical, radular, and morphological characters to produce a cladogram for 26 extinct and extant fissurellid taxa. She recognized a primitive clade comprising F. peruviana and the two extralimital northern hemisphere species (F. nimbosa, F. volcano), a polytomous clade that included South American and South African species with well developed ribs, and a clade with smooth-shelled, weakly ribbed, and finely ribbed taxa. The ensuing description of new morphological groups of Fissurella (Fissurella) is informed by a consideration of species’ temporal ranges but is based largely on shell sculpture. These new groups are compared with clades derived from an analysis of molecular data by Olivares (2006). Groups of related Fissurella (Fissurella) species Coarsely ribbed group: Fissurella maxima, F. costata, and F. picta have long been considered closely related because of their similar coarse radial sculpture (Pilsbry, 1890; McLean, 1984a; Stuber, 1991). All three species occur in Pleistocene deposits of Chile (Herm, 1969); F. picta is also found on Pleistocene marine terraces in Argentina (Aguirre et al., 2005). Of the three taxa, the only Peruvian species, F. maxima, occurs in the upper part of Unit HI at Huacllaco (Figure 2), but not in Fissurella-bearing beds of Unit II or the lower part of Unit III, suggesting that the species may have first appeared following the early late Pliocene. Other coarsely ribbed species (FL concolor, F. aranea) extend the record of this group back to the early Pliocene. ‘imbata’ lineage: Specimens of Fissurella limbata have low, broad, and sometimes obsolete ribs and few The Veliger, Vol) 50;3NoyZ SCALE Contour interval is 50 m. Figure 62. Type localities of Fisswrella mcleani, sp. nov. (DV 1635-1) and F. geoglypha, sp. nov. (Yauca roadcut). secondary ribs. The oldest examples are found in upper Unit II beds at Huacllaco. Some Huacllaco specimens have ribs nearly as well differentiated as those on specimens of late Pliocene F. maxima (compare Figure 28 with Figure 8), suggesting that F. limbata diverged from the coarsely ribbed group during the late early Pliocene. Finely ribbed group: The fossil record of F. latimarginata (late Pliocene to Recent) and F. persica (early Pliocene) shows that finely ribbed species have long constituted a distinct morphological group. Fissurella peruviana, another species with fine ribs, appeared first during the early late Pliocene (Figure 3), suggesting it is more deeply rooted within the group than F. /atimarginata. Less deeply rooted may be F. cumingi, an extant species greatly resembling F. latimarginata that was found on an early middle Pleistocene marine terrace. Smooth-shelled group: A single specimen of Fissur- ella pulchra (Figure 49) was found in upper Pliocene deposits of southern Peru. The discovery of two new late early Pliocene smooth-shelled species, F. mcleani (Figure 55) and F. geoglypha (Figure 59), shows that this group was well established as early as the other morphological groups. ‘crassa’ lineage: The fimbriate wrinkle-ribbed Fissur- ella crassa occurs in early late Pliocene deposits of southern Peru (Figure 2), as well as Pliocene beds in Chile (McLean, 1984a). Its unusual blunt anterior and weakly developed and widely spaced primary ribs crossing an otherwise smooth surface makes its assignment to other groups problematic. Unassigned taxa: Some species of Fissurella (Fissur- ella) are difficult to assign to an existing morphological group. Fissurella radiosa Lesson, 1831, a late Pleisto- cene to Recent species from southern Chile and Argentina, might be placed with finely ribbed taxa, although its narrow ribs are sharply elevated. The T. J. DeVries, 2007 extant F. bridgesii Reeve, 1849, from Peru and Chile, is irregularly striated and ribbed and might belong with finely ribbed taxa. F. nigra Lesson, 1831, a large extant species from southern Chile, has fine striations and is not clearly a member of any group. To summarize: data from western South America point to the existence of at least five fissurellid groups with fossil and modern constituents (Figure 63): a coarsely ribbed group (Ff aranea, F. concolor, F. maxima, F. picta, F. costata), a ‘limbata’ lineage (F. limbata), a finely ribbed group (F- persica, F. peruviana, F. latimarginata, F. cumingi, F. oriens Sowerby, 1835, F-. bridgesii), a ‘crassa’ lineage (F. crassa), and a smooth- shelled group (F. mcleani, F. geoglypha, F. pulchra). The differentiated ribs on some fossil specimens of F. limbata suggest the ‘limbata’ lineage and the coarsely ribbed group (F. maxima and others) share a common early Pliocene ancestor. All morphological groups were already established by the late early Pliocene, implying their common ancestor probably existed no more recently than the earliest Pliocene. To date, however, Fissurella of such antiquity have not been found in southern Peru. Comparison with Molecular Data Olivares (2006) sequenced partial nucleotide se- quences of several mitochondrial and nuclear genes from all 13 of McLean’s (1984a) extant South American species of Fissurella (Fissurella), as well as the extant Caribbean F. nimbosa and extant eastern Atlantic F. schrammi and Diodora graeca (Linnaeus, 1758). His molecular phylogeny, which is remarkably congruent with the phylogeny hinted at by the paleontological data, includes: a. acoarsely ribbed clade that includes F. maxima + F. limbata, b. a deeply rooted clade with finely ribbed taxa, including F. latimarginata + F. cumingi, and a sister group composed of F. pulchra + F. radiosa, c. adeeply rooted F. peruviana that is a sister taxon to other finely ribbed species, and d. a deeply rooted F. crassa with uncertain clade affinities. The most significant incongruency between the molecular and morphological species groupings is the placement of Fissurella radiosa. That the broad, low, smooth-shelled F. pulchra and elongate, high-crowned, sharply ribbed F. radiosa could be sister species (Olivares, 2006) is startling. Yet, every tree-producing algorithm presented Olivares with the same pulchra + radiosa group, no matter which DNA sequence was used. In the case of randomly amplified polymorphic DNA (RAPD) data analyzed using the UPGMA method (unweighted pair group method with arithme- Page 143 3 ~ Ss 3 8 . ~~ ~ . “~~ 80 = OSS TS S SNS ess SS es S dS 5 2 Q Ss 2's ec Sm io S SQ ks as — SS 2 4 SS 62 SS 8 SSG Ts 4 iS © S = Q N as) £6 8 SN wy Q = 3) RECENT LATE PLEIST- PLIOCENE OCENE EARLY PLIOCENE Figure 63. Estimates of temporal ranges for species of Fissurella (Fissurella) found in southern Peru. tic mean), however, F. radiosa was grouped with F. crassa and F. nigra in a deeply rooted clade (Olivares, 2006). The discrepant phylogenetic results for F. radiosa remain unexplained. Molecular ages for branching within the Fissurella (Fissurella) clade of Olivares (2006) are consistently younger compared with ages inferred from paleonto- logical data (Table I). The discrepancies may be due in part to an age estimate for the uppermost terrace in southern Peru (2 Ma) that is too old or molecular calibration points that are too young (e.g., Pleistocene ages for the first appearance of F. maxima and F. peruviana). Diversity Patterns The species richness of Chilean and Peruvian fissurellid faunas for the Pliocene and Quaternary is summarized in Table II. Species richness appears to have increased during the Quaternary, a trend contrary to that for the rest of the Peruvian molluscan fauna (DeVries, 1995, 1997, 2001, 2003). Alternatively, fissurellids may simply be poorly preserved in older deposits. Extinct species of Fissurella are found only in Pliocene beds, not in Pleistocene deposits, in agreement with diversity patterns for the entire Pliocene-Quater- nary molluscan fauna of western South America, which suffered a major species-level extinction during the late Pliocene (Herm, 1969: DeVries, 1985, 2001; Rivade- neira & Marquet, 2007). Large species of Fissurella (>80 mm maximum length) appear only in Quaternary deposits (Table ITI). In contrast, the oldest species of Fissurella (F. aranea, F. persica) are among the smallest species (maximum length < 40 mm). Page 144 The Veliger; Vol. 50; INowZ Origin of South American Fissurella Fissurella (Fissurella) is notable for its taxonomic diversity in western South America since the early Pliocene and complete lack of a fossil record before- hand, even though upper Miocene littoral deposits are common in southern Peru (DeVries, 1998). Miocene Fissurellidae have been identified in Chile (Tavera, 1979: Nielsen, 2003), but the two new taxa illustrated by Nielsen (2003) are probably examples of Diodora Gray, 1821. The holotype of Fissurella alternula Tavera, 1979, might also be a Diodora (Tavera, 1979, 1991); other early Miocene species of Diodora are known from Chile (Nielsen, 2003) and an early Pliocene specimen has been found in southern Peru (Figure 61). South American Fissurella (Fissurella) most likely did not originate from early Miocene Chilean limpets. If Fissurella was introduced to ‘western South America, the avenues for introduction were few: immigration from (1) the Caribbean Sea or eastern North Pacific Ocean by way of northwestern South America, (2) the western Pacific Ocean or Indian Ocean by means of the high latitude West Wind Drift or equatorial countercurrents, or (3) the South Atlantic Ocean. Plausible examples exist for the first two scenarios, e.g., (1) the muricid, Pterorytis Conrad, 1862, arriving in southern Peru from the North Pacific Ocean during the early late Pliocene (DeVries, 2005c), and (2) the turbinid ancestor of Prisogaster Morch, 1850, arriving from the western Pacific Ocean during the middle to late Miocene (DeVries, 2006). No persuasive examples of the third scenario are known (Nielsen, personal communication, 2007). The occurrence of Fissurella peruviana in the Taime Formation of northern Peru (DeVries, 1986, 1988) could be construed as evidence for a _ Pliocene connection with northern hemisphere fissurellids (F: nimbosa, F. volcano) involving either emigration of Caribbean fissurellids southward (Olivares, 2006) or Peruvian taxa northward (Stuber, 1991). The anoma- lous presence of other higher latitude species in the Table I Ages for cladogenetic events affecting Fissurella (Fissurella). See Olivares (2006) for molecular data and basis for estimating ages. Paleontological ages are estimates based on faunal correlations with a radiometrically dated section at Sacaco (Muizon & DeVries, 1985) that also yielded biostratigraphically useful diatoms (H. Schrader, written communication, 1986) and a 2-Ma age for the uppermost uplifted marine terrace in southern Peru. Molecular age Event (Ma) Paleontological age (Ma) Basal polytomy 3.5—2.14 >4 for Fissurella (Fissurella) Divergence of F. limbata and ancestor of F. maxima Fissurella latimarginata species complex polytomy, including divergence of F. latimarginata and F. cumingi 2.18—1.3 >2.3 Taime Formation, however, suggests that a northward expansion of the Peruvian Faunal Province during the late Pliocene (DeVries, 1986) would best explain the odd occurrence of F. peruviana near the Equator. No other species of Fissurella (Fissurella) have been found in Miocene or Pliocene beds in northern Peru or Ecuador (e.g., Olsson, 1932, 1964; Pilsbry & Olsson, 1941; Marks, 1951; DeVries, 1986). Absent such a record, passage of diverse Fissurella populations during the Pliocene through northwestern South America from the Caribbean seems unlikely. The emigration of Fissurella from Peru to California, however, resulting Table II Species numbers of western South American Fissurella (Fissurella) for different time intervals since the early Pliocene. Data from Herm (1969), McLean (1984) and this report. Time interval Recent Species list for chile, peru Number of species bridgesii, costata, crassa, cumingi, latimarginata, limbata, 13 maxima, nigra, oriens, peruviana, picta, pulchra, radiosa Pleistocene peruviana, picta, pulchra Late late Pliocene Early late Pliocene Early Pliocene costata, crassa, cumingti, latimarginata, limbata, maxima, 9 crassa, latimarginata, limbata, maxima, peruviana, pulchra 6 concolor, crassa, limbata, persica, peruviana aranea, concolor, geoglypha, mcleani, persica 5 Nn T. J. DeVries, 2007 Page 145 Table III Maximum of sizes of Fissurella (Fissurella) species through time. Data from McLean (1984) and fossils described in this paper. Large species (maximum length > 80 mm) are shaded black; medium sized species (maximum length 40— 80 mm) are shaded medium gray; and small species (maximum length < 40 mm) are shaded light gray. aranea concolor mcleani geoglypha peruviana S 7) 3 q iS) persica Recent Pleistocene Late late Pliocene Early late Pliocene Early Pliocene in the Pleistocene establishment (Grant & Gale, 1931) of a single species, F. volcano, as proposed by Stuber (1991), cannot be ruled out. The first species of Fissurella (Fissurella) probably arrived in Chile or Peru during the late Miocene or earliest Pliocene, since the subgenus was fully diversi- fied by the late early Pliocene. In southern Peru, late Miocene antecedents to the modern Peruvian mollus- can fauna were already established following a major species-level extinction event between 14 and 11 Ma (DeVries, 2001, 2002; DeVries & Frassinetti, 2003). Late Miocene rocky intertidal and mixed rock-and- sand subtidal molluscan faunas included numerous muricid genera and teguline trochids, as well as barnacles and the inarticulate brachiopod, Discinisca Dall, 1871 (DeVries, 1995, 1997, 2003, 2005a, 2005b). To this mix were added the turbinid, Prisogaster (about 10 Ma), the trochinine trochid, Piscoacritia DeVries and Hess, 2004 (about 7 Ma), and a thaid, Purpura boliviana Philippi, 1887 (about 7 Ma), all probably with western Pacific ancestry (DeVries and Hess, 2004; DeVries, 2006; DeVries, unpublished data). Fissurellids from the western Pacific Ocean or Indian Ocean could have arrived similarly. Other keyhole limpets (Fissur- ellidea group of genera) do show a disjunct South African and southern South American distribution (McLean, 1984b), which lends credence to a Southern Ocean origin for South American Fissurella (Fissur- ella). The difficulty with a western Pacific or South African origin for South American Fissurella is that no obvious candidate for an ancestor exists. Mono- dilepas monilifera (Hutton, 1873), a New Zealand limpet with an early Miocene to Recent record (Dell, limbata latimarginata pulchra maxima cumingi costata picta bridgesii oriens radiosa nigra 1953; Beu & Maxwell, 1990), has a very broad foramen, cancellate sculpture, and lacks an aragonite/calcite shell structure. The same is true for some South African fissurellids [Fissurellidea aperta (Sowerby, 1825)]. Other South African fissurellids, including the coarsely ribbed modern Fissurella mutabilis Sowerby, 1834, and finely ribbed F. natalensis Krauss, 1848, as well as the smooth-shelled fossil F. robusta Sowerby, 1889, and F. glarea Carrington & Kensley, 1969 [late Miocene to late Pliocene (Roberts & Brink 2002; Franceschini & Compton 2004)], share some characters with South America Fissurella, but lack the aragonitic / calcitic shell layers. CONCLUSIONS Fissurella (Fissurella) are two-layered limpets (McLean, 1984a) from intertidal and shallow subtidal habitats along the coast of western and southern South America that may well constitute a monophyletic clade (Olivares, 2006). Fissurella appeared abruptly on the shores of western South America by the late early Pliocene, already fully diversified into coarsely ribbed, finely ribbed, and smooth-shelled groups that are largely congruent with molecularly defined clades (Olivares, 2006). The nearly nonexistent fossil record from northern Peru and examples of Neogene immigration by western Pacific taxa to the shores of western South America favor a Southern Ocean origin for Fissurella (Fissurella), although candidates for an ancestral fissur- ellid taxon are not obvious; late Miocene South African F. robusta and F. glarea might serve best as sister taxa. Fissurella limpets are ecologically important mem- bers of rocky intertidal and subtidal communities in Page 146 Peru and Chile and are abundant enough to be the target of an artisanal harvest (Castilla & Fernandez, 1998). Their sudden appearance in the region during the late Miocene or early Pliocene is likely to have had an impact on trophic dynamics in communities where medium-sized and large limpets were mostly unknown [one exception: Cellana fuenzalida (Herm, 1969), a giant nacellid limpet from Chile and southern Peru (Herm, 1969; Lindberg and Hickman, 1986; DeVries, unpublished data]. The interplay between recruitment of larval Fissurella, barnacles, and competition for space with a variety of algal species, for example, spelled out in the case of modern F. picta by Lopez et al. (1999), would have been repeated for an increasingly more diverse fissurellid fauna during the late early to late Pliocene, a time of global climate cooling (Ravelo et al., 2004) and mass extinction of marine mollusks in the Pliocene Peruvian Faunal Province (DeVries, 2001); opportunities for occupying new and abandoned niches may have expanded at that time. Acknowledgments. I would like to thank Lindsey Groves (Natural History Museum of Los Angeles County, California) for the loan of comparative material from Peru, Chile, South Africa, and New Zealand; L. Groves, James McLean (also of the Natural History Museum of Los Angeles County), Javier Quinteiro (Universidad de Santiago de Compostela, Spain), Alberto Olivares (Universidad de Antofagasta, Chile), Sven Nielsen (Universitat Kiel, Germany), and Robyn Stuber (United States Environmental Protection Agency, California) for their advice and help with finding relevant literature; and Mario Urbina and Marcelo Stucchi (Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru) for advice and logistical help. At various times, this research has benefited from grants from the U.S. National Science Foundation (EAR-85-03886), the Conchologists of America, Inc. (1993), and the Fulbnght Senior Scholarship program (1999). LITERATURE CITED ALAMO, V. & V. VALDIVIESO. 1997. Lista sistematica de moluscos marinos del Peru. Instituto del Mar del Peru: Callao, Peru, 183 pp. AGUIRRE, M. L., S. RICHIANO & Y. NEGRO SIRCH. 2005. 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Gobierno de Chile: Santiago, 256 pp. PitsBRY, H. A. 1890. Stomatellidae, Scissurellidae, Pleuroto- mariidae, Haliotidae, Scutellinidae, Addisoniidae, Coccu- linidae, Fissurellidae. Manual of Conchology 12:323. PitsBRY, H. A. & A. A. OLSSON. 1941. A Pliocene fauna from western Ecuador. Proceedings of the Academy of Natural Sciences of Philadelphia 93:79. RAVELO, A. C., D. H. ANDREASON, M. LYLE, A. OLIVAREZ LYLE & M. W. WARA. 2004. Regional climate shifts caused by gradual global cooling in the Pliocene epoch. Nature 429:263-267. REEVE, L. 1849. Monograph of the genus Fissurella. Conchologia Iconica 6 pls:1-—8. RIVADENEIRA, M. M. & P. A. MARQUET. 2007. Selective extinction of late Neogene bivalves on the temperate Pacific coast of South America. Paleobiology 33(3):455468. ROBERTS, D. L. & J. S. BRINKS. 2002. Dating and correlation of Neogene coastal deposits in the Western Cape (South Africa): implications for neotectonism. Journal of South African Geology 105:337-352. SOWERBY, G. B. 1835a. Shells collected by Mr. Cuming on the western coast of South America and among the islands of the South Pacific Ocean. Proceedings of the Zoological Society of London for 1834:123-128. SOWERBY, G. B. 1835b. A catalogue of the Recent species of Fissurellidae, genus Fissurella. Thesaurus conchyliorum, or Monographs of genera of shells: London, pp. 1-8, 77 figs. STUBER, R. A. 1991. Reconstructing the historical biogeogra- phy of Fissurella (Fissurella) Bruguiere, 1789: a phyloge- netic approach. Masters thesis, University of California: Berkeley, 84 pp. TAVERA, J., J. 1979. Estratigrafia y paleontologia de la Formacion Navidad, Provincia de Colchagua, Chile (Lat. 30°50’S). Boletin del Museo Nacional de Historia Natural, Chile 36:176. TAVERA, J., J. 1991. Contribucion al estudio de la fauna de la Formacion Navidad, revisiones taxonomicas. Chile Cen- tral, Navidad (estratotipos) Lat. 30°51’, Arauco Lat. Page 148 41°40’. Unpublished Report, Departamento de Geologia, Universidad de Chile, Santiago, 476 pp. APPENDIX List of Locality-Samples. ‘GPS’ signifies field measure- ment with GPS position. Yauca La Mina Lomas DV 239-11 DV 401-1 DV 465-1 DV 599-2 DV 810-1 DV 923 Panamerican Highway, roadcut descending towards village and river valley. Late Early Pliocene. Beach at La Mina, Paracas Peninsula. 13°54'32"S, 76°18'58"W (Pisco 1:100,000 quad- rangle). Recent. Intertidal rocks, beach wrack, and trash near port village of Lomas. 15°34’S, 74°49’W (Yauca 1:100,000 quadrangle). Recent. 4.5 km south-southeast of Los Organos, along the Los Organos-Somatito road, cliffs near Occidental Petroleum wells 10531, 10529, and 10536. Lower 25 m of Taime Formation. Estimated coordinates 4°13’S, 81°06’30"W. Late Pliocene. Hueco La Zorra, beach. 14°02'31"S, 76°15'51”"W (Punta Grande 1:100,000 quadrangle). Recent. Marine terrace north of Chala; elevation about 160-170 m above sea level. Middle Pleistocene. Montemar, Pleistocene marine terraces. 15°32’05"S, 74°47'39"W (Yauca 1:100,000 quadrangle). Middle Pleistocene. Quebrada Champeque, about 15 km north of Chala, marine terrace at 200 m above sea level. Fisswrella specimens from surface of terrace; estimated to have been dropped by humans. Holocene. Knolls on southeast side of Rio Acari. 15°36'29"S, 74°37'53"W (GPS; Yauca 1:100,000 quadrangle). Early to late Pliocene. (See Figure 3.) Samples from measured section include DV 923-1 and DV 923-le (both about 9m) and DV 923-1la (0.2 m). DV 1032 DV 1141-1 DV 1251-1 DV 1254 DV 1372-1 DV 1418-1 DV 1628-6 DV 1635-2 LACM 75-31 LACM 75-32 The Veliger, Vol. 50, No. 2 First curves at top of Huacllaco section. 15°52’59"S, 74°10'05"W (GPS; Chala 1:100,000 quadrangle). Includes samples DV 1032-1 (first curve in highway heading south, about 60-70 m in measured Huacllaco section, latest Pliocene) and DV 1032-2 (about 40-45 m in measured Huacllaco section, early late Phocene). See Figure 2. Northwest side Hueco La Zorra, beach. 14°02'46"S, 76°15'58”W (GPS; Punta Grande 1:100,000 quadrangle). Recent. Quebrada Champeque, roadcut along Pana- merican Highway. 15°48'42”S, 74°21'24"W (GPS; Chala 1:100,000 quadrangle). Late Pliocene. Roadcuts along Panamerican Highway 10 km southeast of Chala, above Playa Huacllaco. 15°53’S, 74°09'W (GPS; Chala 1:100,000 quadrangle). Samples from measured section (see Figure 2) and elsewhere include: DV 1254-2 (36.5 m), DV 1254-5 (44.2 m), DV 1254-6 (41.7 m), DV 1254-10 (48 m), DV 1254-14 (41 m), DV 1254-Bal5 (27 m), DV 1254-Ball0 (47.5 m), lowest cobbles (37 m). Late early Pliocene to late Pliocene. See Figure 2. Punta Lomas, intertidal rocks and beaches. 15°34’'S, 74°49'W (Yauca 1:100,000 quadrangle). Recent. South side Acari depression, upper beds. 15°34’50"S, 74°36'59”"W (GPS; Yauca 1:100,000 quadrangle). Huacllaco section southeast of Chala, sandstones of upper Unit III (see Figure 2). Chala 1:100,000 quadrangle. Late Pliocene. Yauca Depression, western side Panamerican Highway. 15°39'33”S, 75°34'54”"W (GPS; Yauca 1:100,000 quadrangle). Late early Pliocene. Intertidal zone and beach, Islota Concon, north of Vina del Mar, Chile. 32°52’S, 71°33’W. Recent. Collected by J. H. McLean, 1975. Intertidal zone, Punta El Lacho, north of Cartagena, Chile. 33°30’S, 71°39’W. Recent. Collected by J. H. McLean, 1975. The Veliger 50(2):149—162 (June 20, 2008) THE VELIGER © CMS, Inc., 2007 Revision of the Protobranch Species Described by Dautzenberg & Fischer (1897) with Description of a New Species and Taxonomic Comments on Bathyspinula (Bivalvia, Nuculanoidea) RAFAEL LA PERNA Dipartimento di Geologia e Geofisica, Universita di Bari, Via Orabona 4, I-70125 Bari, Italy (e-mail: r.laperna@geo.uniba.it) Abstract. In 1897, Philippe Dautzenberg and Henry Fischer described six deep water protobranchs from the North Atlantic (Princesse-Alice expeditions, 1894, 1896): Leda excisa (Philippi) var. subexcisa, Leda bernardi, Leda allaudi, Leda mirmidina, Leda mabillei and Malletia perrieri. Almost all of these taxa are only known from the original description, with no further records in the modern literature. The present revision, based on the original material, led to the following combinations: Bathyspinula subexcisa, Ledella bernardi, Yoldiella allaudi (lectotype designated), Microgloma mirmidina, Nuculana mabillei and Tindaria perrieri. A new species is described as Yoldiella dautzenbergi from material misidentified as Leda allaudi. Taxonomic comments are given for the genus Bathyspinula Filatova, 1958. The subfamily Bathyspinulinae Coan & Scott, 1997 (= Spinulinae Allen & Sanders, 1982 nom. inval.), formerly in the family Nuculanidae, is raised to full family rank within the Nuculanoidea. Tindariopsis Verrill & Bush, 1897 is also assigned to the Bathyspinulidae. INTRODUCTION Dautzenberg & Fischer (1897) described six new deep water protobranchs from the North Atlantic (Prin- cesse-Alice expeditions 1894, 1896): Leda excisa (Phi- lippi) var. subexcisa, Leda bernardi, Leda mabillei, Leda allaudi, Leda mirmidina and Malletia perrieri. Almost all of these taxa are only known from the original description, with no further records in the modern literature on the North Atlantic molluscs. A single species, Spinula subexcisa (Dautzenberg & Fischer, 1897), was included in two taxonomic works (Clarke, 1961; Allen & Sanders, 1982), but without examination of the type material. The present work offers a systematic revision of these poorly known species, based on the original material. This paper also gives an occasion to discuss the taxonomy of the genus Bath- yspinula and its systematic position within the Nucu- lanoidea. MATERIAL AND METHODS The Princesse-Alice stations from which Dautzenberg & Fischer (1897) described the species dealt with in the present work are reported in Figure 1. Dautzenberg (1927) renumbered all the stations from the Princesse- Alice, Hirondelle and Prince de Monaco expeditions (1886-1913) as a single series, with longitudes west of Greenwich, whereas the original longitudes were west of Paris. In the present work, station numbers and longitudes are according to Dautzenberg’s (1927) list, with the original station number in parenthesis. Dautzenberg (1927) also reported the same descriptions and illustrations as those originally published by Dautzenberg & Fischer (1897). The type material is from the Musée Oceanographi- que de Monaco, the Institut Royal des Sciences Naturelles de Belgique, Bruxelles and the Monterosato collection, Museo Civico di Zoologia, Rome. A list of this material is reported in Table 1. The nuculanid classification adopted in the present work follows the scheme by Ockelmann & Warén (1998), except for the position of the genus Bathyspinula. The following abbreviations are used: exp(s). — expedition(s), sh(s) — complete shell(s), paired valves; v(s) — valve(s); IRScN — Institut Royal des Sciences Naturelles de Belgique, Bruxelles; MOM — Musée Oceanographique de Monaco, MCZ — Museum of Comparative Zoology, Harvard University, Cam- bridge, MZR — Museo Civico di Zoologia, Rome. SYSTEMATICS Family Nuculanidae H. & A. Adams, 1858 Genus Nuculana Link, 1807 Nuculana mabillei (Dautzenberg & Fischer, 1897) (Figures 2a—e) Leda mabillei Dautzenberg & Fischer, 1897:207, pl. 6, figs 9, 10. Page 150 The Veliger, Vol. 50, No. 2 5 ae T B » Corvo Flores 85703 Ie fe) 698 Graciosa sd Sao Jorge = ’ Terceira Faial @ — Ov Tt Gy 3,49 Pico ig é Sao Miguel - AZORES e = 7380 in Santa Maria » 37 200 km 30 25 a ee ees ee | | | EEE | Figure 1. Leda mabillei — Dautzenberg, 1927:291, pl. 8, figs 25, 26. Types: Monaco exps., st. 503 (Princesse-Alice 1894, st. 101), 47°10'N, 5°47'45"W, 748-1262 m, 1 v, MOM 21160, holotype. Distribution: Only known from a single deep water station off the Bay of Biscay. Remarks: The single, poorly preserved type right valve is notably robust and convex, triangular-elongate in shape, with a short, truncate, bicarinate rostrum and a wide, slightly concave postero-dorsal area. The sculp- ture consists of commarginal ridges, somewhat irregu- lar in spacing and strength, slightly coarser posteriorly (Figures 2a, d). The hinge is relatively strong, with a triangular, oblique ligament pit (Figure 2e). A shallow, poorly defined rostral ridge is present internally. The rostrum tip is slightly broken (Figure 2d), giving appearance of an oblique truncation, as in the original description (rostrum oblique truncatum). Leda mabillei can be easily assigned to the genus Nuculana. It is different in many respects from the two well known North Atlantic species of Nuculana, i. e. N. pernula (O.F. Miller, 1776) and N. minuta (O.F. Miller, 1776). Good illustrations of these two species were reported by Schiotte & Warén (1992). Due to the short rostrum, Nuculana mabillei is more similar to N. Table 1 List of the type material. MOM IRScN MZR Leda excisa var. subexcisa 5 vs 2 vs Leda bernardi lv 2 vs Leda mabillei lv Leda allaudi lv lv Leda mirmidina 2 vs 16 vs Malletia perrieri lv Monaco expeditions, stations 503, 698, 703, 738, 1114, 1349. minuta, from which it differs mainly by being less elongate and more triangular in shape, more robust and convex. The occurrence of N. mabillei in a deep water station is puzzling, as species of this genus typically occur in shallow waters. Moreover, this species is notably different from some deep water nuculanids with a long, slender and bent rostrum which can be assigned to Thestyleda Iredale, 1929 (Di Geronimo & La Perna, 1997). The valve of N. mabillei could have underwent a down-slope transport from outer shelf bottoms, as also suggested by its poor preservation status. Genus Ledella Verrill & Bush, 1897 Ledella bernardi (Dautzenberg & Fischer, 1897) (Figures 3a-f ) Leda bernardi Dautzenberg & Fischer, 1897:206, pl. 6, figs 5, 6. Leda bernardi — Dautzenberg, 1927:289, pl. 8, figs 21, 2D. Nuculana bernardi — Clarke, 1962:52. Types: Monaco exps., st. 738 (Princesse-Alice 1896, st. 109), 37°40’N, 26°25'15”"W, 1919 m, 1 v, MOM 21158, holotype. Same station as holotype, 2 vs, IRScN 1238/5. Distribution: Only known from a single deep water station, west of Sao Miguel, Azores. Remarks: Leda bernardi was described from a single right valve (MOM) (Figures 3a—c), but two other valves from the same station as the holotype are present at IRScN (Figures 3d-f): one of them is fairly well preserved and younger, the other is badly preserved. The shell is ovate-elongate, not particularly convex, moderately robust, shortly rostrate, with a very shallow subrostral sinuation and an obscure posterior keel. The umbo is strongly opisthogyrate. The surface bears growth striae and ill defined, widely spaced commar- Rese ererna, 2007 Page 151 Figure 2. Nuculana mabillei (Dautzenberg & Fischer, 1897). a-e. Holotype (Dautzenberg & Fischer, 1897:pl. 6, figs 9, 10), length 10.35 mm, MOM 21160. Figure 3. Ledella bernardi (Dautzenberg & Fischer, 1897). a—c. Holotype (Dautzenberg & Fischer, 1897:pl. 6, figs 5, 6), length 9.40 mm, scale bar = 1 mm, MOM 21158. d, e. topotype, length 6.11 mm, IRScN 1238. f. topotype, length 8.20 mm, IRScN 1238/5. Page 152 ginal ridges, becoming better defined near the ventral margin. The hinge is relatively strong, with a triangu- lar, oblique ligament pit. The pallial sinus is well defined, not particularly deep. The larval shell is worn in all three valves. This species shows remarkable similarities with a group of deep-water North Atlantic species which includes Leda oxira Dall, 1927, Leda semen Smith, 1885, Ledella parva Verrill & Bush, 1898 and Ledella similis Allen & Hannah, 1989. They share an ovate- elongate shell, a short rostrum, a poorly defined subrostral sinuation and a strongly opisthogyrate umbo. Of these, Ledella similis is the only species known from the West European Basin (Allen & Hannah, 1989), whereas Leda oxira, L. semen and Ledella parva are from the Western Atlantic (Smith, 1885; Verrill & Bush, 1898; Dall, 1927; Allen & Hannah, 1989). All of them were referred to Ledella by Allen & Hannah (1989) who also attempted to synonymise Ledella parva with Leda semen (see also Verrill & Bush, 1898). This problem is hard to resolve, since the type material of Leda semen is destroyed (Allen & Hannah, 1989), but the examination of the original illustrations (Smith, 1885:pl. 19, figs 2, 2a; Verrill & Bush, 1898:pl. 81, fig. 1) suggests a distinct status for both species. Another deep water species, Ledella librata Dell, 1952 from the Challenger Plateau, New Zealand, seems notably similar to the group of Atlantic species. The systematic position of these species is not clear. They actually recall Ledella, but differ by being notably elongate and with a strongly opisthogyrate umbo. The species of Lede/la usually have a rather sharp posterior keel, a well defined subrostral sinuation and a pointed rostrum (e.g., Warén, 1978; Allen & Hannah, 1989; La Perna et al., 2004). However, no other genus so far described seems to provide a better position for this group of species. Genus Yoldiella Verrill & Bush, 1897 Yoldiella allaudi (Dautzenberg & Fischer, 1897) (Figures 4a—d) Leda allaudi Dautzenberg & Fischer, 1897:207, pl. 6, figs 7, 8. Leda allaudi — Dautzenberg, 1927:290, pl. 8, figs 23, 24. Nuculana allaudi — Clarke, 1962:52. Types: Monaco exps., st. 703 (Princesse-Alice 1896, st. 74), 39°21'20"N, 31°06'W, 1360 m, 1 v, MOM 21156, pe: 1 v, IRScN, 1238/03, paralectotype. : Only known from a single, deep water sta t of Flores, Azores. Other records (Daut- the VeliserVolas50) Now zenberg & Fischer, 1897; Dautzenberg, 1927) cannot be confirmed. Remarks: The shell of Yoldiella allaudi is delicate, markedly convex, ovate, distinctly inequilateral in shape and with a sculpture of thin, irregularly spaced commarginal ridges. The hinge is thin, with anterior and posterior rows of teeth of similar length, separated by a small, triangular ligament pit. The larval shell is ovate, 170 um in maximum length. The material of Leda allaudi at MOM consists of a single valve (st. 703), illustrated by Dautzenberg & Fischer (1897:pl. 6, figs 7, 8). Three valves from the same station are present at IRScN: one of them is Leda allaudi, whereas the two other valves belong to a new species herein described. Additional material labelled as Leda allaudi, from two other stations, is present at IRScN: Monaco exps., st. 1349, 38°35’30"N, 28°05'45”W, 1250 m, Azores and st. 1114, 33°59'30"N, 8°12'45”, 851 m, off Casablanca (Figure 1), but no specimen proves to be Leda allaudi. The material from the Azores includes 3 valves of an unidentified species, here kept as Yoldiella sp. A, and two poorly preserved, unidentifiable shells. The mate- rial from off Casablanca includes Yoldiella semistriata (Jeffreys, 1879) (2 vs, 1 sh), Yoldiella seguenzae Bonfitto & Sabelli, 1995 (2 vs) and an unidentified species (1 v), here kept as Yoldiella sp. B. Some of this material is illustrated in Figure 5. In order to fix the identity of Leda mabillei, a lectotype was designated (Figures 4a, b): it is the left valve from st. 703 (MOM), illustrated by Dautzenberg & Fischer (1897). The other valve from the same station at IRScN is a paralectotype (Figures 4c, d). None of the many species of Yo/diella known from the Atlantic (e.g., Warén, 1989; Allen et al., 1995; Salas, 1996) seems particularly similar to Y. allaudi, except for Y. subaequilatera (Jeffreys, 1879) and the following new species, as discussed below. Yoldiella dautzenbergi n. sp. (Figures 4e—h) Type material: Holotype and one paratype (left valves), IRSecN, 1238/03. Type locality: Monaco exps., st. 703 (Princesse-Alice 1896, st. 74), 39°21’20"N, 31°06’W, 1360 m. Etymology: Named after Philippe Dautzenberg, Bel- gian malacologist (1849-1935). Description: Shell small, thin walled, ovate, poorly elongate, subequilateral, moderately convex. Umbo at mid line, small, slightly opisthogyrate, distinctly protruding from shell outline. Posterior end well rounded, anterior end obscurely rostrate, slightly Rec Perma; 2007 Page 153 Figure 4. a—d. Yoldiella allaudi (Dautzenberg & Fischer, 1897). a, b Lectotype (Dautzenberg & Fischer, 1897:pl. 6, figs 7, 8), length 5.16, MOM 21156. c, d. Paralectotype, length 4.01 mm, IRScN 1238/03. e-h Yoldiella dautzenbergi n. sp. e, f. Holotype, length 4.02 mm, IRScN 1238/03. g, h. Paratype, length 4.08 mm, IRScN 1238/03. narrower than anterior. Ventral margin moderately convex, with a faint slope break at postero-ventral transition. Subrostral sinuation almost absent. Sculp- ture consisting of growth striae and well incised, irregularly spaced commarginal lines, giving appear- ance of wide, flat ribs. Hinge plate thin, delicate. Dentition taxodont, with chevron-shaped teeth in two series of similar length, with about ten teeth anteriorly and posteriorly. Anterior row slightly convex, posterior row almost straight. Ligament pit small, triangular, sunken. Adductor muscle scars ovate, of similar size. Pallial sinus narrow, moderately deep. Prodissoconch ovate, 280 um in maximum diameter. Measurements: holotype 4.02 mm in length, 2.97 mm in height, 1.05 mm in width; paratype 4.08 x 2.90 < 0.95 mm. Distribution: Only known from a single, deep water station, east-of Flores, Azores. Remarks: The material of Yoldiella dautzenbergi n. sp. is from the lot of Leda allaudi at IRScN (see under Yoldiella allaudi). Yoldiella dautzenbergi 1s much less convex, slightly less elongate and more equilateral than Y. a/laudi, with the posterior end just slightly narrower than the anterior one. The umbo is less opisthogyrate and slightly smaller, the ventral margin less convex. In both species there is a faint slope break at the postero- ventral transition, but it is more distinct in Y. dautzenbergi. The subrostral sinuation is almost absent in Y. dautzenbergi and the sculpture consists of deeply incised, irregularly spaced commarginal lines, rather than of thin ridges. The larval shell is smaller than in Y. allaudi. The largest valve of Y. allaudi is about 5 mm in length, whereas the two valves of Y. dautzenbergi are about 4 mm, but the material is too scant for assessing a size difference between the two species. A close resemblance also exists with Yoldiella Page 154 The Veliger, Vol. 50, No. 2 Figure 5. a,b. Yoldiella semistriata (Jeffreys, 1879). Monaco expeditions (1901), st. 1114, off Casablanca, length 3.56 mm, IRScN 1238/03. c, d. Yoldiella seguenzae Bonfitto & Sabelli, 1995. Monaco expeditions (1901), st. 1114, off Casablanca, length 2.58 mm, IRScN 1238/03. e, f. Yoldiella sp. A, Monaco expeditions (1902), st. 1349, Azores, length 3.62 mm, IRScN 1238/03. g, h. Yoldiella sp. B. Monaco expeditions (1901), st. 1114, off Casablanca, length 4.86 mm, IRScN 1238/03. subaequilatera (Jeffreys, 1879), a poorly known deep water species from the Northeast Atlantic, dealt with by Waren (1989:p. 235, figs 10a, b). The new species is slightly less elongate and less equilateral than Y. subaequilatera, and more convex, with a narrower umbonal angle and with a better defined sculpture. Genus Microgloma Sanders & Allen, 1973 Vicrogloma mirmidina (Dautzenberg & Fischer, 1897) (Figures 6a—g) o Dautzenberg & Fischer, 1897:208, pl. Leda mirmidina — Dautzenberg, 1927:292, pl. 8, figs 27— 30. Nuculana mirmidina — Clarke, 1962:53. Types: Monaco exps., st. 698 (Princesse-Alice exp. 1986, st. 69), 1846 m, 39°11’N, 30°44’40"W, 2 vs, MOM 21157, syntypes; 16 vs, MIRScN 1239/02, syntypes. Distribution: Only known from a single, deep water station, south-east of Flores, Azores. Remarks: The position of the family Pristiglomidae Sanders & Allen, 1973 in the superfamily Nuculoidea, as proposed by Sanders & Allen (1973), was criticized by Ockelmann & Warén (1988) who assigned Pristi- R. L. Perna, 2007 Page 155 Figure 6. a-g. Microgloma mirmidina (Dautzenberg & Fischer, 1897). a, b. Syntype (Dautzenberg & Fischer, 1897:pl. 6, figs 12, 14), length 1.84 mm, IRScN 1239/02 (ligament pit enlarged by breaking or corrosion). c. Syntype, length 1.47 mm, IRScN 1239/02. d, e. Syntype, length 1.58 mm, IRScN 1239/02. f, g. Syntype, length 1.63 mm, scale bar =0.5 mm, IRScN 1239/02. h, i. Syntype, length 1.57 mm, MOM 21157. gloma Dall, 1900 and Microgloma Sanders & Allen, 1973 to the Nuculanidae. They also presented strong evidence for the progenetic character of Microgloma. Three further species of Microgloma were known, all from the Atlantic (Sanders & Allen, 1973; Ockelmann & Warén, 1998): M. yongei Sanders & Allen, 1973 (type species), M. tumidula (Monterosato, 1880) (= ™. turnerae Sanders & Allen, 1973) and M. pusilla (Jeffreys, 1879). The last two species occur in European waters. Another European species, Phaseolus guilonardi Hoeksema, 1983 is provisionally placed in Microgloma, but it clearly belongs to a different group, as discussed by Ockelmann & Warén (1998) and La Perna (2003). Microgloma mirmidina is somewhat similar to M. yongei and M. tumidula in the ovate-subrectangular shape, whereas M. pusilla is distinctly egg-shaped. All these species have a comparatively robust, notably convex shell, with a sculpture of thin ridges near the ventral margin. The muscle scars are slightly buttressed in M. mirmidina and generally well-defined in the other species. The largest syntype is 1.84 mm in shell length (Figures 6a, b), the others 1.5-1.6 mm. Microgloma mirmidina therefore is notably larger than M. yongei, M. tumidula and M. pusilla which are about 1 mm in shell length (Allen & Sanders, 1973; Ockelmann & Warén, 1998). The shell shape changes notably with growth, from dorso-posteroventrally oblique to poste- riorly elongate, whereas the other species grow almost isometrically and equilaterally, as seen in the growth series of M. yongei and M. tumidula reported by Sanders & Allen (1973). This is probably due to the relatively large size of M. mirmidina, allowing this species to follow a growth pattern more similar to that of normal sized bivalves, whereas the other species are too small for manifesting marked allometric changes. At a size larger than 1.3-1.5 mm, the growth of M. mirmidina produces a stepped shell edge, giving a box- Page 156 like appearance. As observed by Ockelmann & Warén (1998), such a growth pattern which at a smaller extent occurs in the other species of Microgloma, provides an increase in shell volume and counterbalances the effects of miniaturization. Besides the small size, Ockelmann & Warén (1998) remarked two other synapomorphies for the Micro- gloma species: the enlarged innermost teeth of the left valve and the radially wrinkled surface of the prodissoconch. The first character is not present in M. mirmidina (Figure 6g), but admittedly it is not always present or clearly developed in the other species (e.g., Ockelmann & Warén, 1998:fig. 9f). However, the hinge of M. mirmidina is similar to that of the congeners, with slightly chevron-shaped to rather stout teeth and a small, elongate ligament pit. The ligament pit is slightly oblique, with the anterior end apparently external or semi-external (Figure 6g). It is similar to the oblique ligament pit of a juvenile specimen of Yoldiella Dhilippiana (Nyst, 1845) illustrated by Ockelmann & Waren (1998:fig. 3b), which differs by being posteriorly external. This supports the hypothesis by Ockelmann & Warén (1998:11) for the progenetic ongin of Micro- gloma from Yoldiella or Ledella. The larval shell of MM. mirmidina is ovate, about 180 um in length, notably smaller than that of M. yongei (290 um) and M. tumidula (260-270 um), more similar to that of M. pusilla (195-218 um), according to the data by Sanders & Allen (1973) and Ockelmann & Warén (1998). Under optical magnification the prodis- soconch surface shows an unresolved sculpture and it was not possible to ascertain if it corresponds to the radially wrinkled pattern reported by Ockelmann & Waren (1998). Family Bathyspinulidae Coan & Scott, 1997 Genus Bathyspinula Filatova, 1958 Filatova & Shileyko (1984) pointed out the preoc- cupied status of Spinula Dall, 1908 by Spinula Herrich- Schaeffer, 1856 (Lepidoptera). They replaced the genus name Spinula with Bathyspinula Filatova, 1958, for- merly subgenus of Spinula, and erected the new subgenus Acutispinula. Accordingly, Bathyspinula in- cludes the subgenera Bathyspinula (Bathyspinula) and B. (Acutispinula). Species of the latter differ by a finer, almost absent sculpture and a longer, sharper rostrum (Allen & Sanders, 1982; Filatova & Shileyko, 1984; Coan et al., 2000). The type species are Bathyspinula (B.) oceanica (Filatova, 1958) and Bathyspinula (Acutispinula) calcar (Dall, 1908), respectively. Allen & Sanders (1982) erected the monogeneric subfamily Spinulinae (invalidly based on an junior homonym, replaced with Bathyspinulinae by Coan & Scott, 1997) in the family Nuculanidae to contain the genus Bathyspinula, whereas Filatova & Shileyko The Veliger; Vol) 505 Now (1984) included this genus in the subfamily Ledellinae, family Ledellidae Allen & Sanders, 1982. Ockelmann & Waren (1998) kept the Nuculanidae as a single, undivided family; a systematic view markedly different from the multi-taxa classification by Allen & Sanders (1986). However, as discussed below, there are good reasons for keeping Bathyspinula in a separate position, at a full family rank. The adults of Bathyspinula posses a long, mainly external, amphidetic ligament, with a small internal component (Allen & Sanders, 1982; Di Geronimo & La Perna, 1996) (Figures 7a, b; see also the good illustrations by Knudsen, 1970). The internal ligament tends to a semi-external position and part of it can be seen externally, between the umbones of closed valves (Figure 7b). This condition is more evident in the juvenile stages, which possess a proportionally larger, clearly semi-external ligament pit (Figure 7c). The other nuculanids, such as Nuculana, Ledella and Yoldiella have a juvenile, external amphidetic ligament becoming fully internal with growth, as well-document- ed by Ockelmann & Warén (1998), or leaving a small external relict as in Jupiteria (La Perna et al., 2004). The ligament of Bathyspinula is then much more similar to that of the families Malletiidae H. & A. Adams, 1858 (Sanders & Allen, 1985), Tindariidae Verrill & Bush, 1897 (Sanders & Allen, 1977) and Neilonellidae Shileyko, 1989 (Warén, 1989; Allen & Sanders, 1996; La Perna, 2007), all with a well-developed external ligament and a smaller internal component in the adults, than to that of the other nuculanids. None of these families provide a suitable position for Bath- yspinula, for the following reasons: 1) malletiids have a subrectangular, posteriorly truncate or bluntly rostrate, poorly sculptured shell; 2) neilonellids have an ovate, poorly rostrate shell with no trace of subrostral sulcus and postero-ventral sinuation; 3) tindariids have a roundish, not rostrate shell and are asiphonate (Bath- yspinula has well developed, united siphons: Filatova & Shileyko, 1984; Allen & Sanders, 1982). A full family rank is therefore adopted for the Bathyspinulinae Coan & Scott, 1997 (= Spinulinae Allen & Sanders, 1982). The family Bathyspinulidae also provides a suitable position for Tindariopsis Verrill & Bush, 1897, instead of the Tindariidae (Verrill & Bush, 1898), Malletiidae (Dall, 1898; Vokes, 1980; Laghi, 1986) or even Nuculanidae, subfamily Ledellinae (Allen & Sanders, 1996). The type species, Malletia (Tindaria) agathida Dall, 1889 has the same ligament type as Bathyspinula, with a “‘well-marked dorsal ligamental furrow and a small notch or «socket» under the beak” (Verrill & Bush, 1897, 1898; see also Dall, 1898:582). Tindariopsis agathida has a shallow pallial sinus (Dall, 1898; Allen & Sanders, 1996) and cannot be assigned to the Tindar- iidae (which lack a pallial sinus), as suspected by Verrill & Bush (1898). On the other hand, the pointed, keeled IR, IL, leering), AO Figure 7. a-c. Ligament characters of Bathyspinula. a. Bathyspinula subexcisa (Dautzenberg & Fischer, 1897), Challenger exp. (1973), st. 4, length 5.01 mm, MCZ 348787. b. Bathyspinula hilleri (Allen & Sanders, 1982), st. DS23, length 4.93 mm, MCZ 348807. c. Bathyspinula excisa (Phi- lippi, 1844), Archi, southern Calabria, Early-Middle Pleisto- cene, length 3.25 mm, author’s coll. Scale bars: = 1 mm. rostrum and the well-defined subrostral sulcus make Tindariopsis similar to Bathyspinula and markedly different from malletiids and neilonellids, whereas the resemblance with Ledella is due to convergence. A series of good illustrations, though with some misiden- tification, was published by Laghi (1986:pl. 8, figs la— 6c; Nuculana cfr. pusio Philippi of figs la,b is a Tindariopsis species), including the holotype of Tindar- iopsis agathida. Bathyspinula (B.) subexcisa (Dautzenberg & Fischer, 1897) (Figures 7a, 8a—k, r, s) Leda excisa var. subexcisa Dautzenbeg & Fischer, 1897: 205. Leda (Neilo) excisa var. subexcisa — Dautzenberg, 1927:295. Page 157 Spinula subexcisa — Clarke, 1962:52 (?). Spinula subexcisa — Allen & Sanders, 1982:21, figs 22, 8} Dy 3S Types: Monaco exps., st. 698 (Princesse-Alice 1896, st. 69), 39°11’N, 30°44’40"W, 1846 m, Azores, 5 vs, IRScN 1238/01, syntypes; Monterosato coll., 2 vs, MZR 14423, syntypes. Other material examined: Challenger exp. (1973), st. 4, 56°52'N, 10°01'W, 1993 m, Rockall Trough, 4 shs, 1 v, MCZ 348787 (Allen & Sanders, 1982). Chain 106 exp. (1972), st. 318, 50°26.8’-50°27.3'N, 13°19.9’— 13°20.9'W, 2506 m, off West Ireland, 5 shs, MCZ 348785. Distribution: Bathyspinula subexcisa is known from the North Atlantic (West Europe and Azores), in 1846— 2506 m. Remarks: The history of Bathyspinula subexcisa is closely linked to Nucula excisa Philippi, 1844, described from the Plio-Pleistocene of Southern Italy (Philippi, 1844: p. 46, pl. 15, fig. 4; Di Geronimo & La Perna, 1996). According to Allen & Sanders (1982), the records of Malletia excisa by Jeffreys (1876, 1879) and of Leda excisa by Smith (1885) from the North Atlantic could have been based either on Bathyspinula subexcisa or on Bathyspinula hilleri Allen & Sanders, 1982, both occurring in the North Atlantic, the latter with a much wider Atlantic distribution. Also the record of Spinula subexcisa from the South Atlantic by Clarke (1961) was probably based on a different species, possibly Bathyspinula hilleri. He compared his specimens with material from the Jeffreys coll. and found them “identical to M. excisa, as Jeffreys understood it.” This is the first time the type material of Bathyspinula subexcisa is revised. The sole illustrations so far available for this species were the drawings by Allen & Sanders (1982). The most obvious differences from Bathyspinula excisa (Figures 8n—q) lie in the shallower subrostral sinus and in the finer sculpture. Bathyspinula subexcisa also differs by being less convex and more delicate, with a shorter rostrum and a less distinct rostral keel. Allen & Sanders (1982) reported Bathyspinula sub- excisa from a single station (Challenger exp. 1973, st. 4). Some of this material was examined (Figures 8f—i) and it actually matches the type material. Microscopic, anastomosing radiating lines, are present along the subrostral sulcus; they are similar to the microsculpture present in Bathyspinula excisa (Di Geronimo & La Perna, 1996:pl. 2, figs 1, la). This character is not visible in the type material of B. subexcisa, most probably due to the poor preservation. Other speci- mens (Chain 106 exp. 1972, st. 318) differ by having a Page 158 The Veliger, Vol. 50, No. 2 Figure 8. a-k. Bathyspinula subexcisa (Dautzenberg & Fischer, 1897). a, b. Syntype, length 6.81 mm, IRScN 1238/01. c, d. Syntype, length 6.52 mm, IRScN 1238/01. e. Syntype, length 5.47 mm, IRScN 1238/01. f. Challenger exp. (1973), st. 4, length 4.44 mm, MCZ 348787. g, h. Challenger exp. (1973), st. 4, length 5.01 mm, MCZ 348787. 1. Challenger exp. (1973), st. 4, length 3.09 mm, MCZ 348787. j. Chain 106 exp., st. 318, length 4.42 mm, MCZ 348785. k. Chain 106 exp., st. 318, length 5.35 mm, MCZ 348785. 1, m. Bathyspinula hilleri (Allen & Sanders, 1982). 1. St. DS23, length 3,42 mm, MCZ 348807. m. St. DS23, length 4.93 mm, R. L. Perna, 2007 Page 159 less convex ventral margin (Figures 8j, k), but appar- ently without any clear-cut separation from the specimens with a more convex ventral margin. The larval shell of B. subexcisa is ovate, similar in size and shape to that of B. excisa (Di Geronimo & La Perna, 1996:pl. 2, fig. 3), 280-300 um in length, both in the type material and in the material from MCZ. This contrasts with the size of 450 um reported by Allen & Sanders (1982:23): such a difference must be due to a measurement error. Bathyspinula hilleri was described from the Angola Basin and reported from a number of stations through the Atlantic Ocean (Allen & Sanders, 1982). Some material (Figures 8m, t) from the West European Basin (st. DS23, 46°32.8’N, 10°21’W, no data on cruise and station depth, not reported by Allen & Sanders, 1982, tab. 4), matches the original description. It differs from Bathyspinula subexcisa by having a more convex ventral margin, a slightly coarser sculpture, particularly near the ventral margin, and by being slightly more inflated. Bathyspinula excisa is notably common in the Plio- Pleistocene bathyal deposits cropping out in Italy (Di Geronimo & La Perna, 1996, 1997; La Perna, 2003). The finding of a single, fresh valve in the Ibero- Moroccan Gulf (Salas, 1996) seems to bring evidence that small populations are still present in the adjacent Atlantic. The depth range of Bathyspinula excisa was (or 1s) much shallower, from 200-300 m down to some 1000 m at least, than that of B. suwbexcisa and the other congeners, greatly exceeding 1000 m (Knudsen, 1970; Allen & Sanders, 1997; Olabarria, 2005). Family Tindariidae Verrill & Bush, 1897 Genus Tindaria Bellardi, 1875 Tindaria perrieri (Dautzenberg & Fischer, 1897) (Figures 9a—c) Malletia perrieri Dautzenberg & Fischer, 1897:208, pl. 6, figs 15, 16. Malletia perrieri var. curta Locard, 1898:333, pl. 18, figs 20-24. Malletia perrieri— Dautzenberg, 1927:296, pl. 8, figs 19, 20. Types: Monaco exps., st. 698 (Princesse-Alice 1896, st. 69), 39°11’N, 30°44’40"W, 1846 m, 1 v, MOM 21159, _ holotype. — Distribution: Azores (south-east of Flores) and North- west Africa (off Rabat), 1846-2190 m. Remarks: The holotype is a poorly preserved right valve, somewhat robust, ovate in shape, with anterior and posterior ends well rounded and a strongly anterior umbo. Most of the outer surface bears only growth striae and ill-defined commarginal ridges, becoming better defined, sharper and regularly spaced towards the ventral margin. The hinge is moderately strong, arched with a continuous series of teeth. As observed by Dautzenberg & Fischer (1897), there is no ligament pit. A thin, barely visible external ligament furrow is present posteriorly, slightly extending anteriorly. Malletia perrieri var. curta, described by Locard (1898) from the Talisman st. 16, 2190 m, off Rabat, Morocco (the original coordinates were based on the Paris meridian and the corrected version is 34°01'N, 08°32'W; S. Gofas, pers. comm.) is a synonym of Malletia perrieri. This is supported by the close matching of the two descriptions and the almost perfect overlap of the shell outlines. Locard’s var. curta was said to be slightly higher and shorter, but the two original valves (of the same shell, as inferred from the illustrations) are only slightly larger, 9 mm in length, 8 mm in height, than the holotype of Malletia perrieri (7.93 X 6.92 mm), with the same length to height ratio. According to Sanders & Allen (1977), Tindaria Bellardi, 1875 and Pseudotindaria Sanders & Allen, 1977 (currently in the Neilonellidae) cannot be distinguished from each other conchologically (Pseu- dotindaria differs from Tindaria by having siphons). Warén (1989) remarked that Pseudotindaria has an edentulous gap in the hinge, as in the type species Pseudotindaria erebus (Clarke, 1959). This observation seems more useful for distinguishing the two genera than the assumption by Maxwell (1988) that Tindaria lacks a pallial sinus. The type species of Tindaria is T. arata Bellardi, 1875, from the Late Miocene of the Turin area. The examination of the types and of abundant topotypic material of 7. arata confirmed Wareén’s (1989:255, figs 19c, d) observations: 1) the tooth series is continuous (more precisely, there is a short interruption, sometimes poorly defined, much shorter than the edentulous gap in Pseuwdotindaria); 2) the pallial line is feeble, slightly distinct anteriorly, fading posteriorly and, apparently, without sinus. The characters of Malletia perrieri all point to Tindaria (except for the inability to examine the pallial MCZ 348807. n—q. Bathyspinula excisa (Philippi, 1844). n, o. Archi, southern Calabria, Early-Middle Pleistocene, length 6.21 mm, author’s coll. p, q. Archi, southern Calabria, Early-Middle Pleistocene, length 3.25 mm, author’s coll. r, s. Bathyspinula subexcisa. s. Same as Figure k. t. Challenger exp. (1973), st. 4, length 4.70 mm, MCZ 348787. t. Bathyspinula hilleri, same as Figure m. Page 160 The Veliger, Vol. 50, No. 2 Figure 9. a-c. Tindaria perrieri (Dautzenberg & Fischer, 1897), holotype (Dautzenberg & Fischer, 1897:pl. 6, figs 15, 16), length 7.93 mm, MOM 21159. d-f. Tindaria sp., length 5.96 mm, IRScN 1238/4. Scale bars =1 mm. line because of the poor preservation status). None of the Atlantic tindariids (Sanders & Allen, 1977; Warén, 1989) seems particularly similar to Tindaria perrieri. A single right valve labelled as Malletia perrieri is present at IRScN (Figures 9d-f), from the same station as the holotype at MOM. No mention of this valve was made, either by Dautzenberg & Fisher (1897) or by Dautzenberg (1927). It is rather robust, not markedly convex, with a sculpture of only growth striae. The posterior margin is poorly convex or somewhat truncate, with a slope break at the postero-dorsal transition. An external ligament furrow is present posteriorly. The pallial line is somewhat straight posteriorly, with no pallial sinus. It seems to represent an undescribed species, of uncertain systematic posi- tion, provisionally kept as Tindaria sp. Acknowledgments. I am grateful to Thierry Backeljau (Institut Royal des Sciences Naturelles de Belgique, Bruxelles) and Michéle Bruni (Musée Oceanographique de Monaco) for the loan of material from the Princesse-Alice expeditions, to Lionello Tringali (Rome) for his kind collaboration in accessing the Monterosato collection and to Adam J. Baldinger (Museum of Comparative Zoology) for the loan of material from MCZ. Serge Gofas (Universidad de Malaga) R. L. Perna, 2007 Page 161 is acknowledged for having suggested the present study and for help at various stages. The librarians of the Stazione Zoologica di Napoli “Anton Dohrn” are thanked for having kindly fulfilled all my bibliographic requests. A special thank to Arie W. Janssen (Gozo, Malta) and Anders Warén (Naturhistoriska riksmuseet, Stockholm) for their careful and valuable critical reading. Work supported by Fondi di Ricerca d’Ateneo 2006 (responsible R. La Perna). LITERATURE CITED ALLEN, J. A. & F. J. HANNAH. 1986. A reclassification of the recent genera of the subclass Protobranchia (Mollusca: Bivalvia). Jourrnal of Conchology 32:225—249. ALLEN, J. A. & F. J. HANNAH. 1989. Studies on the deep sea Protobranchia: the subfamily Ledellinae (Nuculanidae). The Natural History Museum (London), Bulletin (Zool- ogy) 55:123-171. ALLEN, J. A. & H. L. SANDERS. 1982. Studies on the deep sea Protobranchia: the subfamily Spinulinae (family Nucula- nidae). 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The taxonomy of some North Atlantic species referred to Ledella and Yoldiella (Bivalvia). Sarsia 63:213-119. WAREN, A. 1989. Taxonomic comments on some protobranch bivalves from the Northeastern Atlantic. Sarsia 74:223— 259. Instructions to Authors The Veliger publishes original papers on any aspect of malacology. All authors bear full responsibility for the accuracy and originality of their papers. Presentation Papers should include an Abstract (approximately 5% of the length of the manuscript), Introduction, Materials and Methods, Results, and Discussion. Short Notes should include a one- sentence Abstract. In taxonomic papers, all names of taxa must be accompanied by author and date of publication, and by a full citation in the bibliography. In papers on other subjects and in the non-taxonomic portions of taxonomic papers, author and date of names need not be accompanied by a full citation. All genus and species names should be in italics. 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SMITHSONIAN INSTI TN CoNnTENTS — Continued | 3 9088 01 Pliocene and Pleistocene Fissurella Bruguiére, 1789 (Gastropoda: Fissurellidae) from Southern Peru THOMAS {J UIDE VRIES) Syojciec er lusiec peut eo lance Sensis eu ee Oa G eeeiel eel eases ee Cece ee AE ee ee 129 Revision of the Protobranch Species Described by Dautzenberg & Fischer (1897) with Description of a New Species and Taxonomic Comments on Bathyspinula (Bivalvia, Nuculanoidea) RAPABT TACRER NAG oasis ee sc cut cele eee EERE nso Sete TSN ene TE Se crn ee 149 (Oly Mv OU ISSN 0042-3211 A Quarterly published by = CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. ZS Berkeley, California R. Stohler (1901-2000), Founding Editor ] line ees enett I ABKRy, Volume 50 October 1, 2008 Number 3 CONTENTS Valiguna flava (Heynemann, 1885) from Indonesia and Malaysia: Redescription and Com- parison with Valiguna siamensis (Martens, 1867) (Gastropoda: Soleolifera: Veronicel- lidae) SUZETE RODRIGUES GOMES, JULIANE BENTES PIcANGO, MENNO SCHILTHUIZEN AND JOSE WILLIBALDO THOME Review of the Genera Ividia, Folinella, Oscilla, Pseudoscilla, Tryptichus and Peristichia (Gas- tropoda, Pyramidellidae) from Brazil, with Descriptions of Four New Species ALEXANDRE D. PIMENTA, FRANKLIN N. SANTOS AND RICARDO S. ABSALAO ..........--. 171 Galba truncatula Miiller, 1774 (Pulmonata: Lymnaeidae) in Argentina: Presence and Natural Infection by Fasciola hepatica (Linnaeus, 1758) (Trematoda: Digenea) Laura Issta, Sttvia M. PieETROKOVSKY, FLORENCIA KLEIMAN, PaBLO CARMANCHAHI AND (CRISTINA, WARSINASS ae CODES Biante Gen Giclee oto. a acinioio e ploreta.cecececic mic ci aio oeeea ore ree enced 185 Diel Patterns of Vertical Distribution in Euthecosomatous Pteropods of Hawaiian Waters DANTEIMEMINTIEGROPANDIRO CERURASEADYA Siler o amin a abemiae nnicioe ca ni oes 190 Two New Species of Doriopsilla from the Tropical Western Atlantic with Remarks on Cariop- sillidae Ortea & Espinosa, 2005 SN GEIR ALDES AND JEBEL EIAMANNs105 cl o56 5. sa iGend aes oe od eis pe peeves 210 Dendropoma mejillonensis sp. nov., a New Species of Vermetid (Caenogastropoda) from North- ern Chile ATED OFLACHECOPANDE|ORGEN) WAUDIENS GS qo ecise centers oie elt aiclcha chelsea eteieapel neers cts 29 ConrtTENTS — Continued The Veliger (ISSN 0042-3211) is published quarterly by the California Malacozoological So- ciety, Inc., % Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. 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Send all business correspondence, including subscription orders, membership applications, payments, and changes of address, to: The Veliger, Dr. Henry Chaney, Secretary, Santa Barbara Museum of Natural His- tory, 2559 Puesta del Sol Road, Santa Barbara, CA 93105, USA. Send manuscripts, proofs, books for review, and correspondence regarding editorial matters to: Geerat Vermeij, Department of Geology, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). The Veliger 50(3):163—170 (October 1, 2008) THE VELIGER © CMS, Inc., 2007 Valiguna flava (Heynemann, 1885) from Indonesia and Malaysia: Redescription and Comparison with Valiguna siamensis (Martens, 1867) (Gastropoda: Soleolifera: Veronicellidae) SUZETE RODRIGUES GOMES,' JULIANE BENTES PICANCO,’ MENNO SCHILTHUIZEN?® AND JOSE WILLIBALDO THOME? "Departamento de Zoologia, Instituto de Biociéncias, Universidade Federal do Rio Grande do Sul. Av. Bento Gongalves, 9500, Cep 91.501-970, Porto Alegre, Rio Grande do Sul, Brazil (e-mail: suzetebio@yahoo.com.br) *Faculdade de Biociéncias, Pontificia Universidade Catélica do Rio Grande do Sul. Av. Ipiranga, 6681, Cep 90.619-900, Porto Alegre, Rio Grande do Sul, Brazil (e-mail: jbp_yahoo.com.br) > Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Locked Bag 2073, 88999 Kota Kinabalu, Malaysia and National Museum of Natural History ‘Naturalis’, P.O. Box 9517, 2300 RA Leiden, the Netherlands (e-mail: schilthuizen@naturalis.nl) *Escritorio de Malacologia e de Biofilosofia, Praga Dom Feliciano, 39, sala 1303 (Ed. Guanabara — Centro), 90020-160, Porto Alegre, Rio Grande do Sul, Brazil Abstract. We redescribe and figure Valiguna flava, an almost unknown Southeast-Asian land slug. Detailed morphology, radula, jaw and living specimens of Valiguna flava were studied for the first time, based on material recently collected in Borneo and on the holotype. V/. flava is also compared with Valiguna siamensis, the only other species of the genus, which is for the first time recorded in China. Key Words: Morphology, anatomy, land slug, Borneo, Australasian region. INTRODUCTION Veronicellidae Gray, 1840 includes a large number of species of land slugs totally without shell (Thomé et al., 2006). Currently, 27 generic taxa are recognized for the family, which are distributed widely in the humid tropics and subtropics (Hoffmann, 1925; Forcart, 1953; Thomé, 1975; Gomes & Thome, 2004). For the Oriental and Australian regions, Gomes & Thomé (2004) recognized six different genera (total of 13 species): Filicaulis Simroth, 1913, Laevicaulis Simroth, 1913, Sarasinula Grimpe & Hoffmann, 1924, Semper- ula Grimpe & Hoffmann, 1924, Valiguna Grimpe & Hoffmann, 1925 and Veronicella Blainville, 1817. Originally, the taxon Valiguna was proposed by Grimpe & Hoffmann (1925a) as a subgenus of Semperula to include Vaginula schneideri Simroth, 1895 from Eastern Sumatra (Indrapura, Tandjong Kuba). However, in the same year that Grimpe & Hoffmann (1925a) proposed the taxon Valiguna, they rejected it and considered Vaginula schneideri a subspecies of Semperula siamensis (Martens, 1867) (Hoffmann, 1925; Grimpe & Hoffmann, 1925b). Only later, when Hoffmann (1941) found specimens which he considered a new species very close to Vaginula schneideri (V1. isseli Hoffmann, 1941), he reconsidered Valiguna, this time as a genus. Hoffmann (1941) included in Valiguna, VI. schneideri and V1. isseli, species in which the vas deferens does not open terminally (acrocaulis form, such as species of Sarasi- nula) nor basally in the penis (p/eurocaulis form, such as species of Semperula), having an intermediate form, the acropleurocaulis or pseudopleurocaulis form. Recently, Gomes & Thomé (2004) examined the holotype of Vaginula flava Heynemann, 1885 (consid- ered by Hoffmann, 1925 a synonym of Semperula maculata) and also the original and subsequent descriptions of Vi schneideri and VI. isseli (Simroth, 1895; Grimpe & Hoffmann, 1925a, b; Hoffmann, 1925, 1941). They concluded that these three species are synonyms and that Valiguna flava is the valid name since it has priority. Gomes & Thomé (2004) also proposed the inclusion of Semperula siamensis into the genus Valiguna because this species also has an acropleurocaulis penis. According to Gomes & Thomé (2004), VL flava has records from Nias, Borneo and Indrapura (Indonesia), and Valiguna siamensis from Galle (Sri Lanka) and Petshaburi (Thailand). Both Page 164 species were insufficiently described in the original description. Vaginulus reticulatus Westerlund, 1883, which is a synonym of V/. siamensis according to Gomes & Thome (2004), was redescribed by Thomé (1984), although V/ flava has not been studied since then. Our primary purpose is, for the first time, to describe and figure in detail the morphology, radula, jaw, and living specimens of Valiguna flava. The study is based on samples recently collected from Borneo and on the holotype. The species is also compared with Valiguna siamensis, the only other species of the genus, based on material from China and on the paratypes of Vaginulus reticulatus (synonym of Vi. siamensis). MATERIAL AND METHODS Six living and adult specimens of V/. flava, which were collected in Borneo and deposited in the Museum BORNEENSIS of Universiti Malaysia Sabah, were analyzed. The holotype and paratype of this species, deposited in the Natural History Museum (BMNH- 1880.10.6), London, England were also studied. V1. flava was also compared with four specimens of V1. siamensis deposited in the Museum of the Institute of Zoology, Chinese Academy of Sciences, Beijing (China) (lots ZMIZ01091, ZM1Z01092), from Yunnan and Guangxi, China. The type-material of Vaginulus reticulatus Westerlund, 1883 (13 specimens) (synonym of VI. siamensis according to Gomes & Thome, 2004), deposited at Swedish Museum of Natural History (Stockholm, Sweden) was also examined (lots SMNH 6427, SMNH 3753). The specimens of both species were dissected under a stereomicroscope for studying the internal structures. Drawings were done using a camera lucida. Two jaws and two radulae (from each species) were extracted under the stereomicroscope and later analyzed using a scanning electron microscope (SEM Philips XL 30). The terminology used and the morphological and anatomical characteristics described and illustrated are those usually considered diagnostic in the Veronicelli- dae (Hoffmann, 1925; Coifmann, 1935; Forcart, 1953; Barker, 2001; Thomé et al., 2002; Gomes & Thome, 2001, 2004). Valiguna flava (Heynemann, 1885) (Figures 1—12) Vaginula flava Heynemann, 1885:10—-11. Vaginula schneideri Simroth, 1895:7-8. Semperula ( Valiguna) schneideri, Grimpe & Hoffmann 1925a: 391-392. Semperula siamensis schneideri; Hoffmann, 1925: 181— 182; Grimpe & Hoffmann, 1925b:18—19, 31-33. Valiguna schneideri; Hoffmann, 1941:236. The Veliger, Vol. 50, No. 3 Figures 1-2. dorsal position of a living specimen (BOR/MOL 3411); 2, ventral position of a fixed specimen (BOR/MOL 3439). External characteristics of Valiguna flava. |, Valiguna isseli Hoffmann, 1941:234. Diagnostic Features The main diagnostic structure in Veronicellidae is the penis. The penis of Valiguna flava has a glans and a base completely distinct from each other. The base is a cylindrical structure, without peculiarities in outer surface. The glans starts from a flap that surrounds the distal extremity of the penis base. First, it is cylindrical, but then it curves, forming a peak, in whose extremity the vas deferens opens. In the dorsum of the curvature (penis apex) is a cylindrical structure covered by dentate and serrated formations (Figures 9-12). External Characteristics The specimens are relatively large and they have an oval body (Figures 1—2). The notum (dorsal region) is smooth only with some widely spaced granules. There are some scattered blackish spots and also a narrow light line on the dorsum (Figure 1). This last one is in the middle of the notum and is not always clearly visible. The notum ground coloration ranges from pale S. R. Gomes et al., 2007 Page 165 Ss 6 Od << ; 4% ry ‘ SDD e< SPP >? > DBD >> > W Y-y: Be ve Ww, ¢ aa: 6 wy wy, NG SAE ne wy, Ae RS > oS S SSRSSSSSSSeee > Figures 3-8. Radula and jaw of Valiguna flava (BOR/MOL 3409). 3, entire jaw; 4, lateral (L), central (C) and lateral (L) teeth, respectively; 5. lateral view of a central tooth; 6. medium part of the right half of the radula; 7, lateral view of a lateral tooth; 8, lateral teeth in the edges of the radula. yellowish brown to dark reddish brown. The hypono- tum is also pigmented from pale yellowish brown to dark reddish brown, depending of the notum colora- tion. However, it is always much lighter than the notum and has a homogeneous coloration, without spots or lines or only with few tiny blackish spots (Figure 2). The sole is light beige and very narrow, having less than half of the hyponotum width. The female pore is situated at ca. 45% of the length of the body measured from the front, and it is far from the pedal groove by ca. 2/5 the width of the hyponotum. In all specimens the female pore is surrounded by a slender line of black pigmentation (Figure 2). Holotype measurements (mm): body length (70.00), body width (29.00), sole width (5.44) and hyponotum width (8.11). Measurements (mm) (6 other specimens): body length (45-60), body width (22-30), sole width (2.5-4.8) and hyponotum width (5.2—10.5). Digestive System The mouth is followed by a buccal bulb (= pharynx), where the radula and the jaw lie. There are two salivary glands with very slender, ramified and delicate acini connected to the buccal bulb. The buccal bulb is connected to the esophagus, which is followed by the gastric crop. The latter is barely delimited from the esophagus, both having almost the same diameter. The gastric crop leads into a stomach, which is long (twice longer than wide). Two lobules of the digestive gland open into the stomach, one anterior and another one posterior. The anterior lobe does not totally cover the Page 166 The Veliger, Vol. 50, No. 3 Figures 9-12. Reproductive system in Valiguna flava (BOR/MOL 3409). 9, complete reproductive system; 10-12, three different positions of the penis (cb, bursa copulatrix; db, duct of the bursa copulatrix; dd, distal posterior deferens; df, vas deferens; dp, proximal posterior deferens; fc, fertilization complex; fg, female genital pore; gb, albumen gland; gh, hermaphroditic gland; gn, glans; jd, canalis junctor; It, long tubules, md, middle deferens; ol, spermioduct; ov, oviduct; pk, peak, in whose extremity the deferens opens; pl, penial gland papilla; pr, prostate; rc, rectum; rp, penis retractor muscle; st, short tubules; sv, seminal vesicle; tf, structure covered by dentate and serrated formations; vg, penis base. Scale bar: | mm. S. R. Gomes et al., 2007 Page 167 anterior intestinal loop. The intestine starts from the stomach, follows to the anterior region where it forms a loop (the anterior intestinal loop) and then continues back to the posterior region. Near the height of the female pore, the intestine penetrates in the body wall, where the rectum begins. The rectum continues to the end of the body where it opens via the anus. The latter is located centrally in the body and is totally hidden over the sole. The anal opening is represented by a fissure. Neither a circular opening (with a defined border) nor an opercular membrane 1s found. The anal opening floor is formed by well developed folds that sometimes seem papillae. The nephridium does not have an external aperture. It is probably connected to the rectum within the body wall. The jaws are well arquated and narrow. They are formed by an average of 24 wide and superposed ribs. The three ribs of the middle are somewhat higher and less distinct from the others (Figure 3). The radula has 93—95 teeth per transverse row, which is formed by one symmetrical central or rhachis tooth (Figures 4-5), flanked on both sides by 46-47 lateral teeth (Figures 6-7). The dental formula is C/ 1+L46—47/2. The lateral teeth are smaller towards the edges of the radula (Figure 8). Pedal and Pallial Nerves, Pedal Aortic System One pair of pallial and one of pedal nerves run from the central nervous system towards the posterior extremity of the body cavity (Barker, 2001). They are parallel and together from the central nervous system until near the height of the bursa copulatrix. After, they are parallel but separated from each other and run like that until the end of the body cavity. The pedal aortic artery begins at the level of the central nervous system and runs between the nerves (centrally) until around the level of the bursa copulatrix. Pedal Gland The pedal gland, located on the anterior extremity of the sole under the head, is short and straight. It is broad in its proximal portion (in the aperture), narrowing in the middle and with the posterior extremity rounded and somewhat broadened (produc- ing a goblet-shape). It has two different areas: one external lighter and one internal yellowish. The posterior extremity of the gland receives a wide pedal gland artery (Coifmann, 1935). Reproductive System The bursa copulatrix (= spermatheca or gametolitic gland) is almost circular, but somewhat concave in the middle. It has a cylindrical duct a little longer than the bursa copulatrix. The canalis junctor (Barker, 2001; Gomes & Thome, 2004) penetrates in the duct of the bursa copulatrix, near to the half of the total length of the duct. The oviduct is wider near to the female genital pore, involving the duct of the bursa copulatrix (Figure 9). The penis has a glans and a base well differentiated. The base of the penis is somewhat cylindrical and smooth. The glans starts from an surrounding structure (as a flap) at the distal extremity of the penis base; it is initially cylindrical and smooth, but quickly tapers and curves forming a peak, in whose extremity the vas deferens opens. In the dorsum of the curvature is a cylindrical structure covered by dentate and serrated formations, which is in the distal extremity of the penis (Figures 9-12). The penial gland is small and has a length similar to the length of the penis. It is formed by a papilla and by about 15 short tubules (they exceed a little the height of the pericardium). The papilla is relatively long, with around half of the tubules’ length. The tubules are not differentiated in groups according to length, although subtle differences in tubule lengths is observed. Distribution: Islands of Sumatra and Borneo (Fig. 18). Natural history: They were found on the forest floor, at night and early in the morning, in one case more or less clustered around a rotten log. Type-material: Holotype. INDONESIA, Borneo: P. E. Gerrard col. (BMNH_ 1880.10.6.4). Other materials. MALAYSIA, Sabah: Danum Valley Field Centre, ca. 60 km W of Lahad Datu, primary dipterocarp forest, ca. 300 m asl. (04°58’00"N 117°48'00"E), 2 specimens, 2003.80.W, P. Craze & M. Schilthuizen col. (BOR/ MOL 3409); Danum Valley Borneo Rainforest Lodge, 1 specimen, 2000.84.W, C. Rutjes col. (BOR/MOL 3410); Kota Kinabalu Distr., Poring Hot Springs, 660 m asl. (06°02'87"N 116°42’16"E), 1 specimen, 2001.59.W, M. Schilthuizen & P. Koomen col. (BOR/ MOL 3411); Interior Prov., Tenom (behind Perkasa Hotel), 327-400 m asl. (05°07’17’”N_ 115°56'40”E), 1 specimen, 2003.89.W, M. Schilthuizen col. (BOR/MOL 3438); Kinabalu National Park (Poring Hot Springs), 500 m asl., 1 specimen, 2003/03/24 (2003.26.W), M. Schilthuizen col. (BOR/MOL 3439). The lot deposited at Natural History Museum in London (England) (BMNH_ 1884.1.10.1), which is identified as a paratype of V/. flava, is not V1. flava. In this specimen (adult), that has not been dissected yet, the penis (the main diagnostic characteristic) is completely different. It is probably an unknown species of Veronicellidae. Page 168 The Veliger, Vol. 50, No. 3 Figures 13-17. Reproductive system in Valiguna siamensis (ZM1Z01092). 13, complete reproductive system; 14, detail of the bursa copulatrix; 15—17, three different positions of the penis (cb, bursa copulatrix; db, duct of the bursa copulatrix; dd, distal posterior deferens; df, vas deferens; dp, proximal posterior deferens; fc, fertilization complex; fg, female genital pore; gb, albumen gland; gh, hermaphroditic gland; gn, glans; hc, honeycomb aspect structure; jd, canalis junctor; It, long tubules, md, middle deferens; ol, spermioduct; ov, oviduct; pk, peak, in whose extremity the deferens opens; pl, penial gland papilla; pr, prostate; rc, rectum; rp, penis retractor muscle; st, short tubules; sv, seminal vesicle; vg, penis base). Scale bar: 1 mm. S. R. Gomes et al., 2007 Chin India °. e Valiguna flava + Valiguna siamensis Page 169 Philippines Indonesia i ee S Figure 18. Map showing the distribution of Valiguna flava and Valiguna siamensis, considering bibliography records and the lots recently collected in the north of Borneo and China. DISCUSSION VI. flava and VI. siamensis are clearly close species. They share several morphological characteristics, although important differences are also found between them. The penis is the structure that most notably discriminates V/. flava from VI. siamensis and even from the other species of the family. The dentate and serrated formation found in the distal extremity of the penis of Vi flava is very characteristic and it is found only in this species of Veronicellidae. In V/. siamensis, the vas deferens also opens in a lateral peak and a well developed base is found. But, in this species the penis has a formation like a “honeycomb” located in the distal extremity (Figures 13-17). It is a complex structure, which is practically absent in juvenile specimens. The bursa copulatrix region has also some differences. In V/. siamensis, the bursa copulatrix duct is extended in the medium region, assuming a domed aspect. In V/. siamensis the bursa duct is fairly longer than the copulatrix bursa, different from Vi. flava. In this species, the bursa is small and assumes a form from spherical to elliptical. In both species, the canalis junctor penetrates in the bursa copulatrix duct, not in the bursa itself (as in many other species of Veroni- cellidae). The other internal characteristics are very similar between V/. flava and VI. siamensis. The characteristics of the digestive system as well as radula and jaw do not show significant differences between both species. Also the disposition of the pairs of pedal and pallial nerves and pedal aortic system are the same in V/. flava and VI. siamensis. The pedal gland is also similar. More- over, all the other described characteristics regarding on the reproductive system (as copulatrix bursa region and penial gland) are very similar between both species. The body form differs between V7. flava and VI. siamensis. The first one has an oval body while the other is longer and narrower, although in both species the sole width is smaller than half the width of the right hyponotum. Unfortunately, the coloration can not be compared since the specimens of V/. siamensis from China were all completely discolored. The specimens of Vs. reticulatus examined by Thomé (1984), and again by us, were also discolored. Externally, V/. flava can be identified mainly by presence of wide blackish spots in the notum, although there is some variation in the intensity of the notum ground coloration (Figure 1). VI. siamensis, previously redescribed by Thome (1984) when he studied Va. reticulatus from Galle (synonyms), is recorded for the first time to China (Yunnan and Guangxi) (Figure 18). Acknowledgements. We are grateful to Centro de Microscopia e Microanalises (CEMM) of Pontificia Universidade Catolica do Rio Grande do Sul for the SEM photomicrographs; to Peter Koomen for the photo of the living specimen; Suzete R. Gomes, Juliane Picango and José W. Thomé gratefully acknowledges financial support from “‘Conselho Nacional de Desenvolvimento Cientifico e Tecnologico.”” Menno Schilthui- zen gratefully acknowledges financial support from the Royal Netherlands Embassy, Kuala Lumpur. REFERENCES BARKER, G. M. 2001. Gastropods on land: phylogeny, diversity and adaptive morphology. In: G. M. Barker (ed.), The biology of terrestrial molluscs. CABI Publish- ing: New York. Pp. 1-146. COIFMANN, I. 1935. Il sistema arterioso della Vaginula solea. Bolletino di Zoologia 6(1/2): 118-122. Page 170 FORCART, L. 1953. The Veronicellidae of Africa (Mollusca, Pulmonata). Annales du Musée du Congo Belge, Sciences Zoologiques 23:1—110. GoMEs, S. R. & J. W. THOME. 2001. 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Ein Beitrag zur Kenntnis ihre Biologie, Anatomie, Systematik, geogra- The Veliger, Vol. 50, No. 3 phischen Verbreitung und Phylogenie. Jenaische Zeits- chrift fur Naturwissenschaft 61(1/2):1—374. HOFFMANN, H. 1941. Uber einige Vaginuliden auf Grund bisher fiir verschollen gehaltener Typstiicke. Zoologischer Anzeiger 136:229-242. MARTENS, E. VON. 1867. Die Preussische Expedition nach Ost-Asien. Nach amtlichen Quellen. Zoologischer Theil, II. Band: die Landschnecken. K6niglichen Geheimen OberHofbuchdrickere: Berlin. 447 pp. SIMROTH, H. 1895. Eine neue Vaginula-Species: Vaginula schneideri n. sp. Sitzungsberichte der Naturforschenden Gesellschaft 19/21:7-8. THOME, J. W., S. R. Gomes & J. B. PICANCO. 2006. Os carac0dis e as lesmas dos nossos bosques e jardins. Useb: Porto Alegre. 123 pp. THOME, J. W. 1975. Os géneros da familia Veronicellidae nas Americas (Mollusca, Gastropoda). Iheringia 48:3— 56. THOME, J. W. 1984. Veronicellidae (Mollusca, Gastropoda) pantropicais: III. Redescrigao de 5 espécies, com base no exame de tipos depositados no “‘Naturhistoriska Rikmu- seet,’ de Estocolmo, Suécia. Theringia (64):29-46. THOME, J. W., S. R. Gomes & R. S. SILVA. 2002. Redescription of the genus and species Heterovaginina limayana (Lesson, 1830) (Gastropoda, Soleolifera, Ver- onicellidae). The Nautilus 116:79-88. THE VELIGER ec ae) The Veliger 50(3):171-184 (October 1, 2008) © CMS, Inc., 2007 Review of the Genera /vidia, Folinella, Oscilla, Pseudoscilla, Tryptichus and Peristichia (Gastropoda, Pyramidellidae) from Brazil, with Descriptions of Four New Species ALEXANDRE D. PIMENTA,' FRANKLIN N. SANTOS? AND RICARDO S. ABSALAO?? "Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, Rio de Janeiro, RJ, Brazil, CEP: 20940-040 (e-mail: adpimenta@yahoo.com.br) * Departamento de Zoologia, Instituto de Biologia, Universidade do Estado do Rio de Janeiro, Avenida Sao Francisco Xavier 524, Maracana, Rio de Janeiro, RJ, Brazil, CEP 20550-900 (e-mail: columel@yahoo.com.br) * Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro (e-mail: absalao@hotmail.com) Abstract. The taxonomy of the species belonging to the genera /vidia Dall & Bartsch, 1904, Folinella Dall & Bartsch, 1904, Oscilla A. Adams, 1961, Pseudoscilla Boettger, 1901, Tryptichus Morch, 1875 and Peristichia Dall, 1889 from Brazil is reviewed. The following species are reported: [vidia havanensis (Pilsbry & Aguayo, 1933), Folinella robertsoni (Altena, 1975), Pseudoscilla babylonia (C. B. Adams, 1845), Peristichia agria Dall, 1889, Oscilla somersi (Verrill & Bush, 1900) and Triptychus niveus Morch, 1875; the last two species are for the first time recorded from Brazil. Four new species are described: Triptychus litosbathron n.sp. is characterized by its smooth base; Oscilla notialis n.sp. and Oscilla aquilonia n.sp. differ in the degree of projection of the protoconch nucleus, and both species are closely related to Oscilla tornata (Verrill, 1884), differing in details of spiral sculpture; and Peristichia lepta n.sp. is distinguished from other Perictichia species by its slender shell, absence of a columellar fold, and by the numerous spiral cords on the base. Key Words: Mollusca, Pyramidellidae, Odostomiinae, taxonomy, Brazil, [vidia, Folinella, Oscilla, Pseudoscilla, Tryptichus, Peristichia. INTRODUCTION The family Pyramidellidae from Brazil is the subject of a major taxonomic review that has already yielded some published results (Pimenta et al., 2000; Pimenta & Absalao 2001a, 2001b, 2002, 2004a, 2004b; Absalao et al., 2003), in addition to others in preparation. In each of these papers, selected genera were studied, common- ly resulting in amendments to the taxonomic status of pyramidellid species reported from Brazil, as well as revealing several new species and expanding the geographic distributions for known species in the Western Atlantic. In this paper we deal with the Odostomiinae and Pyramidellinae genera Ividia Dall & Bartsch, 1904, Folinella Dall & Bartsch, 1904, Oscilla A. Adams, 1861 , Pseudoscilla Boettger, 1901, Trypti- chus Morch, 1875 and Peristichia Dall, 1889. ~ The supraspecific classification of the family Pyr- amidellidae is controversial. There is no consensus about the status of most of the more than 300 generic or subgeneric names (Schander et al., 1999). Particu- larly, in most genera of the subfamily Odostomiinae, the characters of the shell overlap somewhat, and clear differences cannot be established. While some authors (e.g., Dall & Bartsch 1904, 1909; Abbott 1974; Diaz & Puyana 1994) consider the genus Odostomia in a very broad sense, with many subgenera, others have been using many of these subgenera at full generic rank, giving rise to narrower definitions of each genus. Many of these genera have been interpreted differently in several studies in different geographic areas (e.g., Robertson 1978; Jong & Coomans 1988; Linden & Eikenboon 1992; Schander 1994; Pefias et al., 1996: Penas & Rolan 1998; Redfern 2001, among others), often giving rise to different generic allocations for the same species. We believe that a consensus will be reached only after more detailed studies, including the careful comparison of type species, and eventually adding anatomical or molecular data. Our goal, in this paper, is not to provide precise definitions for supraspecific taxa, but rather a more taxonomically accurate knowledge of the diversity and geographic range of the Brazilian pyramidellid fauna. In most cases, we adopted a conservative option, Page 172 following previous allocations of the species herein studied, thus avoiding new combinations. It should be clear, then, that most of the generic allocations used herein are to be considered provisional and changes based on new evidence are to be expected in the future. Abbreviations used: —Collections: ANSP - Academy of Natural Sciences of Philadelphia, Philadelphia, USA: Col.Mol.UERJ - Colecao de Moluscos da Universidade do Estado do Rio de Janeiro; CR - collection Colin Redfern; DOUFPE - Departamento de Oceanografia da Universidade Federal de Pernambuco, Recife; IBUFRJ - Instituto de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro; MNHN - Museum National d’Histoire Naturelle, Paris; MNRJ - Museu Nacional / Universidade Federal do Rio de Janeiro, Rio de Janeiro; MZSP - Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo; USNM - National Museum of Natural History, Washington, DC; YPM - Peabody Museum of Natural History / Yale University, New Haven; ZMA - Zodlogisch Museum Amsterdam, Amsterdam; -—Expeditions: AMASSEDS - A Miultidisciplinary Amazon Shelf Sedimentary Study, Research Vessel ““Columbus Isel- ing” Coll.; GEOMAR XII - Comissao Oceanografica Geologia Marinha XII, NOAS Coll.; MD55 — Marion Dufresne Expedition; PADCT - Programa de Apoio ao Desenvolvimento Cientifico e Tecnologico, Research Vessel “‘Professor W. Besnard” coll.; REVIZEE - Recursos Vivos da Zona Economica Exclusiva, Re- search Vessel “‘Professor W. Besnard” coll. (except when noted); —Colectors: AG - Research Vessel “‘Astro Garoupa,” Petrobras SA; Eq. Zoo Coll. - equipe do Departamento de Zoologia, IBUFRJ; NOAC - Re- search Vessel “‘Almirante Camara,” Marinha do Brasil; NOAN - Research Vessel “Antares,” Marinha do Brasil; NOAS - Research Vessel ‘Almirante Sal- danha,” Marinha do Brasil. MATERIAL AND METHODS The determination of the material was based on comparisons against type material and/or original descriptions and illustrations. In the material examined lists, the number inside brackets indicates the number of shells in each lot. This report is based entirely on empty shells from Brazilian and foreign collections. All lots from MNHN were collected along the northeast Brazilian coast by P. Maestrati from 1984 to 1989. SYSTEMATICS Subfamily Odostomiinae Pelseener, 1928 Genus /vidia Dall & Bartsch, 1904 Ividia Dall & Bartsch, 1904: 11. Type species: Parthenia armata Carpenter, 1857, Mazatlan, by original designation. The Veliger, Vol. 50, No. 3 Ividia havanensis (Pilsbry & Aguayo, 1933) (Figure 1A—E) Odostomia (Miralda) havanensis Pilsbry & Aguayo, 1933: 118, pl. 6, fig. 4: Odé & Speers (1972: 9, not illustrated); Abbott (1974: 298, fig. 3627); Vokes & Vokes (1983: 32, pl. 30, fig. 12); Diaz & Puyana (1994: 235, pl. LXIX, fig. 934). Miralda havanensis: Olsson & McGinty (1958: 44, pl. 1, fig. 8); Rios (1970: 134, 1975: 143, pl. 43, fig. 665, 1985: 165, pl. 54, fig. 784, 1994: 188, pl. 62. fig. 877); Jong & Coomans (1988: 124, pl. 6, 26, fig. 651); Mello (1990: 40, fig. 9); Barros (1994a: 74, not illustrated). Ividia havanensis: Redfern (2001: 144, pl. 65, fig. 595). Type material: Holotype ANSP 159722. Type locality: La Chorrera, Habana, Cuba. Material examined: Para state: IBUFRJ 13693, AMAS- SEDS sta 3228, [1]; —Rio Grande do Norte state: DOUFPE 5012, sta 17 (05°01.317’S / 36°23.507'W, 9.1m), OS5/ii1v/2002, [1]; DOUFPE 5013, sta 30 (04°47'30.837"S / 36°40'02.678"W, 17.3 m), 27/vii/ 2002, [1]; —Pernambuco state: IBUFRJ 11153, Ilha Rata, Fernando de Noronha Archipel, 08/vii/1999, [3]; MNHN, Cabo (enseada dos corais), [16]; MNHN, Cabo (praia de Gaibu), [12]; MNHN, Sao Luiz (areia preta) Maranhao, [3]; MNHN, Paulista (Maria Fari- nha), [2]; MNRJ 10822, Paulista (praia da Concei¢ao), [2]; MNHN, Recife (praia do Pina), [2];}; MNHN, environs de Recife, [6]; MNHN, Cabo (Pedras Pretas), [8]; MNHN, Paulista (praia da Conceigao), [1]; — Sergipe state: DOUFPE 5011, sta 9.2 (11°30'08 "S / 37°07'56 "W, 1031 m), 19/iv/2002, [1]; —Bahia state: IBUFRJ 6374, off Abrolhos, 1990, [3]; —Espirito Santo state: IBUFRJ 8573, off Camburi, 1993, [15]; IBUFRJ 8998, off Aracruz, 18/xii/1989, [1]; IBUFRJ 8481, off Piuma, 1993, [1]; IBUFRJ 8656, off Piima, 1993, [1]; — Rio de Janeiro state: Col.Mol.UERJ 1977, Enseada de Dois Rios, Hha Grande, 19—20/xi/1996, [1]; Col.Mo- 1UERJ 3336, Ilha Grande sta 16 (Rochedo Sao Pedro, 23°2.868'S / 44°32.722'W, 10 m), [1]; Col.Mol.UERJ 3349, Ilha Grande sta 15 (Ponta Grande Timuiba, 23°3.762'S / 44°36.038'W, 7 m), [8]; IBUFRJ 13688, praia da Figueira, Angra dos Reis, 1998, [7]; —Sao Paulo state: MZSP 86785, PADCT sta 6577 (25°15.76’S / 45°04.62’W, 124 m), off Sao Paulo state [1]. Distribution: USA: Florida (Abbott 1974), Texas (Ode & Speers 1972); Caribbean: Habana, Cuba (Pilsbry & Aguayo 1933); Yucatan Peninsula, Mexico (Vokes & Vokes 1983); Panama (Olsson & McGinty 1958); Colombia (Diaz & Puyana 1994); Bahamas (Redfern 2001); Aruba, West Indies (Jong & Coomans 1988); Brazil: Atol das Rocas and Alagoas (Rios 1994); A. D. Pimenta et al., 2007 Page 173 Figure 1. A—E, Jvidia havanensis (Pilsbry & Aguayo, 1933): A, holotype (ANSP 159722); B, E, MNRJ 10822; C—D, Col.Mol.UERJ 1977. A—C, whole shells (respective lengths: 1.9 mm; 1.7 mm; 1.7 mm); D, protoconch; E, detail of last whorl; F, holotype of Ividia abbotti (Olsson & McGuinty, 1958) (ANSP 211912, length: 2.0 mm); G-—I, Folinella robertsoni (Altena, 1975) (MNRJ 10823): G, whole shell (length: 1.6 mm); H, last whorl; I, protoconch. Scale bars: 200 Um. Canopus, Ceara (Barros 1994a); Pernambuco (Mello 1990; this study); Para, Rio Grande do Norte, Fernando de Noronha Archipel, Maranhao, Sergipe, Bahia, Espirito Santo, Rio de Janeiro and Sao Paulo (this study). Remarks: The genus /vidia was proposed by Dall & Bartsch (1904) as a subgenus of Odostomia. This taxon was later synonymized under Miralda A. Adams, 1863 by Dall & Bartsch (1909, p. 172), who argued that the type species that they selected for Jvidia (Parthenia armata Carpenter, 1857) should be referred to Miralda. Ividia havanensis (Figure 1A—E), Ividia abbotti (Ols- son & McGuinty, 1958) (Figure 1F) and also Folinella robertsoni (see discussion below) were, in fact, included in Miralda, as a full genus (Olsson & McGinty 1958; Altena 1975; Rios 1994; Jong & Coomans 1988) and as a subgenus of Odostomia (Pilsbry & Aguayo 1933; Odé & Speers 1972; Abbott 1974; Vokes & Vokes 1983; Diaz & Puyana 1994). Although Schander et al. (1999) adopted the synonymy between Miralda and Ividia, Odé (1993), on the other hand, considered that Parthenia armata is not congeneric with the type species of Miralda (Parthenia diadema A. Adams, 1860), and therefore considered Jvidia a valid genus. This position was followed by Turgeon et al. (1998) and Redfern (2001). In fact, the illustration of Miralda diadema provided by Dall & Bartsch (1906, pl. XVII, fig. 2) is quite distinct from the illustration of Ividia armata by Dall & Bartsch (1909: pl. 19, fig. 6). Ividia armata has a conical shell with two strong nodulose spiral cords in each teleoconch whorl, and weaker spiral cords on the base; whereas Miralda diadema has a somewhat globose shell with a pattern of axial ribs and spiral cords forming nodules where they cross. We, therefore, follow Odé (1993) in considering /vidia a valid genus. Considering /vidia valid, Odé (1993) included Jvidia abbotti; Redfern (2001) included Jvidia havanensis and Ividia robertsoni. However, while we agree with Odé (1993) and Redfern (2001) in regard to 1. abbotti and I. havanensis, we propose the new combination Folinella robertsoni (see below). We consider that the shell of the Page 174 The Veliger;, Vol) 50, Noss latter differs considerably from typical [vidia species, in having three spiral cords in each whorl, crossed by thin axial ribs, lacking the nodulose spiral cords (Figure 1G— H), which fits well with the description of Jvidella by Dall & Bartsch (1909: 172-174, pl. 18, figs. 11, 11a). Lia decorata Folin, 1873 was referred to Miralda by Odé & Speers (1972: 9) and may prove to be an additional species of [vidia, because the shell also has the nodulose spiral cords. Faber (1988) stated that “Odostomia havanensis Pilsbry & Aguayo, 1933 =Lia decorata De Folin, 1873,” but provided no further discussion on this possible synonymy. The illustration provided by de Folin (1873: pl. 6, fig. 8) indeed resembles /. havanensis, but has a shell with spiral nodules very close to each other and with more rounded summits. Besides the records from the east coast of the U.S.A. and the Caribbean, Ividia havanensis was listed and illustrated from Brazil by Rios (1994), in the genus Miralda; however, Rios provided no illustrations of specimens from Brazil, but reproduced the original figure. Other records from Brazil include those of Mello (1990) and Barros (1994a), from two localities on the northeast coast. We now present records of this species from nearly the entire Brazilian north, northeast and southeast coast, considerably enlarging its known geographic range in the Western Atlantic, to about 25°S. Genus Folinella Dall & Bartsch, 1904 Folinella Dall & Bartsch, 1904: 10, nom. nov. pro Amoura de Folin, 1873 non Amoura Forbes, 1845. Type species: Amoura anguliferens de Folin, 1873, by monotypy. Folinella robertsoni (Altena, 1975) new combination (Figure 1G—I) Miralda robertsoni: Altena (1975: 75, fig. 30a—b); Mello (1990: 40, fig. 8); Rios (1994: 188, pl. 62. fig. 878); Barros (1994b: 44, fig. 12a). Odostomia (Miralda) cf. robertsoni: Diaz & Puyana (1994: 235, pl. LXIX, fig. 935). Ividia robertsoni: Redfern (2001: 144, pl. 65, fig. 596). Type locality: Shell ridge near Cupido on the Mar- atakka, Nickerie District, Suriname. Material examined: Rio Grande do Norte state: DOUFPE 5015, sta 01 (05°04.260'S / 36°20.303'W, 3.6 m), 06/vili/2002, [2]; —Pernambuco state: MNHN, Paulista (Maria Farinha), [19]; MNHN Cabo (praia de Gaibu), [10]; MNRJ 10823 Paulista (praia da Con- cei¢ao), [5]; MNHN Cabo (enseada dos corais), [8]; MNHN Raposa (praia de Raposa), [15]; MNHN Itamaraca (praia de Jaguaribe), [1]; MNHN Recife (praia de Candeias), [8]; MNHN Recife (praia do Pina), [8]; MNHN Recife (praia do Pina), [11]; — Maranhao state: MNHN Sao Luiz (areia preta), [5]; — Espirito Santo state: IBUFRJ 10856, Baia de Vitoria, 18/xu1/1998, [2];—-Rio de Janeiro state: IBUFRJ 7477, off Arraial do Cabo, 1993, [2]; Col.Mol.UERJ 3340, Ilha Grande sta 15 (Ponta Grande Timuiba, 23°3.762’S / 44°36.038’W, 7m), [2]; Col.Mol.UERJ 3351, Ilha Grande sta 15 (Ponta Grande Timuiba, 23°3.762’S / 44°36.038’W, 7 m), [2]; IBUFRJ 13689, Angra dos Reis (praia da Figueira), 1998, [4]. Distribution: Suriname (Altena 1975): Caribbean: Colombia (Diaz & Puyana 1994); Bahamas (Redfern 2001); Brazil: Pernambuco, Maranhao (Mella 1990; Barros 1994b; this study); Rio Grande do Norte; Espirito Santo; Rio de Janeiro (this study). Remarks: The taxonomy and nomenclature of Folinella were discussed by Aartsen (1984), Aartsen et al. (1998) and Schander et al. (1999), who considered it as a senior synonym of Jvidella Dall & Bartsch, 1909. As demonstrated by Aartsen (1984), the type species of Folinella is Amoura anguliferens. This genus 1s charac- terized by numerous axial and two to three spiral ribs, equally strong and forming small knobs at their crossings; this sculpture invades the base of the shell. The same pattern of sculpture can be found in some of the species listed by Dall & Bartsch (1909) in the genus Ividella, which should thus be referred to Folinella (e.g., F. quinquecincta Carpenter, 1856, F. delmontensis Dall & Bartsch, 1907, F. navisa Dall & Bartsch, 1909, F. araniana Dall & Bartsch, 1909). The species herein included in Folinella was consid- ered to belong in Miralda, as a full genus (Altena, 1975; Mello, 1990; Rios, 1994; Barros, 1994b) or as a subgenus of Odostomia (Diaz & Puyana, 1994). Red- fern (2001), on the other hand, used the combination Ividia robertsoni. The new combination Folinella robertsoni is pro- posed herein based on similarities to the concept of Folinella, as adopted by Aartsen et al. (1998). Folinella robertsoni has three spiral cords in each whorl, crossed by thin axial ribs, with small nodulose spiral cords (Figure 1G—I), which fits well within the concept of Folinella (fide Aartsen et al., 1998). Folinella robertsoni was listed and illustrated by Rios (1994), in the genus Miralda. He provided no illustra- tions of specimens from Brazil, but reproduced the original figure, and recorded the range of the species in Brazil as “northeast.” Other records from northeastern Brazil are those of Mello (1990) and Barros (1994a). The present paper enlarges the known geographic range of F. robertsoni in the Western Atlantic up to about 23° S, on the coast of Rio de Janeiro. Genus Oscilla A. Adams, 1861 A. D. Pimenta et al., 2007 Page 175 Figure 2. A—F, Oscilla notialis n.sp.: A-E, holotype (MNRJ 10692); F, paratype (IBUFRJ 7595); A—B, whole shell (length: 1.9 mm); C, detail of sculpture on last whorl; D, Last whorl; E-F, protoconch; G—L, Oscilla aquilonia n.sp.: holotype (MNRJ 10825): G—-H, whole shell (length: 1.53 mm); I-J, protoconch; K, detail of sculpture on last whorl; L, last whorl; M, holotype of Oscilla tornata (Verrill, 1884) (USNM 30083). Scale bars: 200 um. Oscilla A. Adams, 1861. Type species: Monoptygma (Oscilla) cingulata. Oscilla notialis n.sp. (Figure 2A—P) Type material: Holotype: MNRJ 10692, off Sergipe state, sta 2.1 (11°23’21"S / 37°04'30’”W, 99 m), 16/iv/ 2002. Paratypes: —Sergipe state: IBUFRJ 14077, type locality [3]; DOUFPE 5000, type locality, [9]; DOUFPE 5006, sta 2.3 (11°24'14"S / 37°05'08’W, 99 m), 16/iv/2002, [2]; —Espirito Santo state: IBUFRJ 14082, off Espirito Santo state, REVIZEE Central I sta D1 (20°48'72"S / 41°09'33”W), 23/11/1996, [1]; —Rio de Janeiro state: IBUFRJ 12316, Cabo Frio VII sta 6194, Page 176 off Rio de Janeiro state, 11/1983, [2]; MNHN, Cabo Frio VII sta 6194, off Rio de Janeiro state, 11/1983, [3]; IBUFRJ 9627, Arquipélago de Santana sta 2, Macaé, 3-5/v/1993, AG coll., [1]; IBUFRJ 6934, off Rio de Janeiro state, GEOMAR XII sta 89 (21°47.8'S / 40°16'W), 28/vii/1979, [3]; IBUFRJ 7595, off Rio de Janeiro state, GEOMAR XII sta 76 (21°57’'S / 40°51'W), 28/viii/1979, [2]; MZSP 86791, PADCT sta 6617 (21°51.6'S / 47°42'W, 327 m), off Rio de Janeiro state [5]; -Sao Paulo state: MZSP 89799, REVIZEE sta 6662 (24°00.95'S / 43°55.54'W, 135 m), [11]; MNRJ 10942, REVIZEE sta 6669 (24°7.42'S / 44°42.22'’W, 101 m), [8]; MZSP 86792, PADCT sta 6571 (24°12.74'S / 44°58.98'W, 79 m), [29]; MZSP 86800, REVIZEE sta 6666 (24°17.13'S / 44°12.15’'W, 163 m), [1]; MZSP 86797, PADCT sta 6579 (24°42.302’'S / 45°18.831'W, 84 m), [1]; MZSP 86787, PADCT sta 6573 (24°42.608'S / 44°43.419'W, 155 m), [1]; MZSP 86794, PADCT sta 6577 (25°15.76’S / 45°04.62'W, 124 m), [14]; MZSP 86788, PADCT sta 6541 (26°15’S / 45°53'W, 130 m), [5]; —Parana state: MZSP 86790, PADCT sta 6631 (25°46'S / 45°28.8'W, 164 m), [2]; —Santa Catarina state: MZSP 86798, PADCT sta 6595 (26°23.55’S / 46°39.49'W, 175 m), [9]; MZSP 86801, PADCT sta 6641 (26°15’S / 45°53’W, 130m), [1]; MNHN, REVIZEE sta 6695 (26°17.134'S / 46°41.788'W, 153 m), [3]; MZSP 86793, PADCT sta 6606 (27°48.07’S / 47°24.04'W, 175 m), [1]; MZSP 86795, PADCT sta 6635 (27°10.38'S / 47°27.54'W, 129 m), [2]. Type locality: off Sergipe state coast (11°23’21"S / 37°04'30"W, 99 m). Distribution: Only known from Brazil. Northeast coast: Rio Grande do Norte and Sergipe states; southeast and south coasts: Espirito Santo, Rio de Janeiro, Sao Paulo, Parana and Santa Catarina states. Etymology: From the Latin noftialis: southern. Refer- ring to the main distribution area of this species, in southern latitudes off Brazil. Diagnosis: Oscilla species with planispiral hetero- strophic protoconch; strongly conical shell bearing wide spiral cords and grooves in teleoconch whorls, and an additional, narrower spiral cord above the suture, variably expressed. Description: Shell conic, holotype 1.9 mm in length. Teleoconch with up to 5 whorls with crenulated outline. Suture deep. Protoconch heterostrophic pla- nispiral, with about 1.5 whorls, smooth, forming an angle of about 90° to shell main axis, diameter about 200 um. Spiral sculpture formed by three cords: the widest cord just above the suture; a second cord of medium width in the middle of each whorl; and a third, narrower cord just above the suture, partially covered by the subsequent whorl, almost inconspicuous in early The Veliger, Vol. 50, No. 3 whorls; between the cords, there are two wide spiral furrows with microscopic axial growth lines. Base slightly convex, with microscopic axial ribs and with its adapical periphery marked by a peripheral, smooth spiral cord that corresponds to the third spiral cord in the teleoconch whorls, and bordered adapically by two additional, very thin spiral cords; with a chink-like umbilicus. Aperture rhomboid, with a columellar fold somewhat projected. Outer lip thin and slightly crenulated. Oscilla aquilonia n.sp. (Figure 2G—L) Type material: Holotype: MNRJ 10825, off Para state, AMASSEDS sta 3228 (03°25.1'N, 49°56.4’W, 64 m), 17/v/1990, RVCI coll. Paratypes: IBUFRJ 4169, type locality, [1]; IBUFRJ 14895, type locality, [3]; MNHN, type locality, [2]; MNRJ 10819, type locality, [3]; IBUFRJ 14897, off Para state, AMASSEDS sta 4134, RVCI coll., [1]. Type locality: off Para state, north Brazilian coast - AMASSEDS sta 3228 (03°25.1'N, 49°56.4’W, 64 m). Distribution: Only known from Para state, north Brazil. Etymology: From the Latin aquilonius: north. Referring to the occurrence of this species on the northern coast of Brazil. Diagnosis: Oscil/la species with helicoid heterostrophic protoconch with strongly projected nucleus; conical shell bearing wide spiral cords and grooves in teleoconch whorls, and an additional, narrower spiral cord above the suture, variably expressed. Description: Shell conic, holotype 1.14 mm in length. Teleoconch with up to three whorls, with crenulated outline. Suture deep. Protoconch heterostrophic heli- coid, with about 1.5 whorls, well projected, smooth, forming an angle of about 90° to shell main axis, diameter about 200 um. Spiral sculpture formed by three cords: the widest cord just above the suture; a second cord, of medium width in the middle of each whorl; and a third, narrower cord, just above the suture, partially covered by the subsequent whorl, almost inconspicuous in early whorls; between the cords, there are two wide spiral furrows with micro- scopic axial growth lines. Base slightly convex, with microscopic axial ribs and with its adapical periphery marked by a peripheral, smooth spiral cord that corresponds to the third spiral cord in the teleoconch whorls; with a small chink-like umbilicus. Aperture rhomboid, with a columellar fold, medium projected. Outer lip thin and slightly crenulated. A. D. Pimenta et al., 2007 Page 177 Figure 3. A-C, Oscilla somersi (Verrill & Bush, 1900) (IBUFRJ 13667): A—B, whole shells (respective lengths: 1.5 mm; 1.6 mm); C: protoconch; D-E, Pseudoscilla babylonia (C. B. Adams, 1845) (IBUFRJ 8490): D, whole shell (length: 1.6 mm); E, protoconch. Scale bars: 200 um. Oscilla somersi (Verrill & Bush, 1900) new combination (Figure 3A—C) Odostomia (Evalea) somersi Verrill & Bush, 1900: 533, pl. 65, fig. 7. Miralda havanensis auct non (Pilsbry & Aguayo, 1933): Sa et al. (1984: 7, fig. 41). Menestho somersi: Jong & Coomans (1988: 124, pl. 6, fig. 652); Redfern (2001: 144, pl. 65, fig. 598A; pl. pl. 113, fig. 598B). Boonea somersi: ? Odé (1993: 55, not illustrated). Boonea someri (sic): Wise (2001: 74, not illustrated). Type locality: Bermuda. Material examined: Bahamas Islands: CR 3641, 26°39'N, 77°18'W, 0.3 m, 20/vii/1982 [5]; CR 10098, NE of Chub Rocks (26°44’N, 77°12'W, 23 m), 12.vii.1992 [1]; -West Indies: ZMA, Aruba, F. Verbene coll. [28]; ZMA, Curagao, Jong coll. [2]; —Brazil: — Espirito Santo state: IBUFRJ 8689, off Piuma, [36]; — Rio de Janeiro state: IBUFRJ 13667, Praia da Figueira, Angra dos Reis, C. Alvarenga coll. [14]; MNHN, Praia da Figueira, Angra dos Reis, C. Alvarenga coll. [6]; MNRJ 10820, Praia da Figueira, Angra dos Reis, C. Alvarenga coll. [6]; coll. Mol. UERJ 5961, Ponta do Pinto, Enseada de Parati-Mirim (23°13.249’S / 44°38.950'W, 5 m), Ilha Grande Bay, [12]; coll. Mol. UERJ 6213, Ponta Grande Timuiba (23°3.762’S / 44°36.038’W, 9 m), Ilha Grande Bay, [22]; coll. Mol. UERJ 6244, Ponta Escalvada, Ilha da Gipoia (23°1.911'S / 44°22.734'W, 11m), Ilha Grande Bay, [35]; coll. Mol. UERJ 6234, Ilha Comprida, Baia de Ribeira (23°57.724'S / 44°22’W, 9 m), Ilha Grande Bay [17]; coll. Mol. UERJ 6256, Laje do Coronel (23°5.884’S / 44°24,410’'W, 23 m), Ilha Grande Bay [15]. Distribution: USA: Texas (Odé 1993): Caribbean: Bermuda (Verrill & Bush 1900); Bahamas (Redfern 2001); Curagao, West Indies (Jong & Coomans 1988); Brazil: southeast coast (this study). Remarks: The genus Oscilla was proposed as subgenus of Monoptygma by A. Adams (1861) and diagnosed with elevate spiral cords, aperture subquadrate, with a columelar fold. Tryon (1886) used Oscilla as a section in the Pyramidella subgenus Syrnola. Van Aartsen (1994) used Oscilla as a full genus, recording Oscilla jocosa from Mediterranean. Schander et al. (1999) considered Oscilla as a genus in the subfamily Chrysallidinae. The three species herein included in this genus fit well such diagnosis, with a conical shell, strong spiral cords and a distinctly visible columellar fold. Other pyramidellid genera have a similar pattern of strong spiral cords, such as Pseudoscilla, Cingulina and Menestho. Oscilla somersi (new combination here proposed), for example, has already being allocated to the genus Menestho by Jong & Coomans (1988) and Redfern (2001). However, Menestho is a_ genus originally described based on Turbo albulus Fabricius, 1780, a species from Greenland. Also, Oscilla somersi was included in Boonea by Ode (1993) and Wise (2001), a position with which we do not agree, since Boonea has three nodulose spiral cords in each whorl. The original figure of Oscilla somersi shows smooth spiral cords. In the shells from Brazil and the West Indies that we examined, we could find some variation in the first two spiral cords, which range from almost completely smooth to sculptured with small nodules of Page 178 varying strength. The shell illustrated by Redfern (2001) also bears somewhat nodulose spiral cords, but some variation in the spiral ridges also can be found (Redfern, personal communication). Odé (1993), al- though expressing doubts about his generic allocation, found variation in the spiral ridges too. According to Johnson (1989), the shells in a lot labeled as syntypes (YPM 15710) do not correspond to this species. Oscilla notialis (Figure 2A—F) and Oscilla aquilonia (Figure 2G—L) differ from the holotype of Oscilla tornata Verrill, 1884 (new combination, herein pro- posed), from off Cape Hatteras, U.S.A. (Figure 2M), in respect to the very narrow upper spiral cord and wider spiral grooves between the cords. Furthermore, Verrill (1884) did not report an additional, narrower spiral cord above the suture, which is found in Oscilla notialis (Figure 2A—B) and also in very similar speci- mens from Abaco (Bahamas), named Oscilla sp. B by Redfern (2001: pl. 65, fig. 600), which may prove to be this species too. This additional spiral cord is a variable character, being stronger in some shells and absent in others. In Oscilla aquilonia, an additional, suprasutural cord may also be present, to a variable degree (Figure 2G, E). Oscilla aquilonia is very similar to Oscilla notialis, but has a helicoid protoconch (Figure 2I-J), whereas Menestho notialis has a planispiral one (Figure 2E—F). Furthermore, O. aqulinonius does not have the two thin spiral cords at the periphery of the last whorl and abapically adjacent to the suture, as seen in O. notialis (Figure 2C_—D). Oscilla notialis shows a wide range of variation, both in shell size (up to five teleoconch whorls) and ornamentation (spiral cords sometimes with subqua- drate edges, and umbilicus wider in some shells). Oscilla aquilonia, on the other hand, is consistently smaller, with almost no variation in the above characters. In spite of the variation found in M. notialis, the protoconch is always planispiral (Fig- ure 2E—F). This is the first record of Oscilla somersi from the southwestern Atlantic, where it has been collected off the southeast Brazilian coast (about 23°S). Oscilla notialis has a wide range of distribution in Brazil, from the northeast (about 4°S) to the southeast coast (about 20°S). Oscilla aquilonia, on the other hand, was found only in northern localities (about 03°N). Genus Pseudoscilla Boettger, 1901 Pseudoscilla Boettger, 1901. Type species: Oscilla (Pseudoscilla) miocaenica Béettger, 1901. Pseudoscilla babylonia (C. B. Adams, 1845) (Figure 3D—E) Chemnitzia babylonia C. B. Adams, 1845: 6: Odé & The Veliger; Voly 50) Noms Speers (1972: 6, not illustrated); Clench & Turner (1950: 259, not illustrated) Chemnitzia (Miralda) babylonia: Morch (1875: 165, not illustrated). Odostomia (Cingulina) Babylonica: Tryon (1886: 358, not illustrated). Chemnitzia Babylonia: Bush (1899: 176, not illustrated). Odostomia (Cingulina) Babylonia: Verril & Bush (1900: 534, pl. LXV, fig. 11). Menestho babylonia: Odé & Speers (1972: 8, not illustrated). Odostomia (Miralda) judithae Newell-Usticke (1959: 86): synonymized by Jong & Coomans (1988). Cingulina babylonia: Warmke & Abbott (1962: 148, not illustrated); Abbott (1974: 301, not illustrated); Rios (1985: 165, pl. 54, fig. 785, 1994: 188, pl. 62, fig. 876); Vokes & Vokes (1983: 32, pl. 30, fig. 19); Jong & Coomans (1988: 120, pl. 19, fig. 637); Mello (1990: 41, fig. 10). Cingula babylonia: Oliveira (1992: 285, not illustrated). Pseudoscilla babylonia: Odé (1993: 58, not illustrated). Odostomia babylonia: Wise (1996: 445, figs. 13a—e); Redfern (2001: 142, pl. 64, fig. 585). Type material: lost (Clench & Turner 1950) Type locality: Jamaica. Material examined: Pernambuco state: MNHN, Cabo (praia de Gaibu), [1]; —Bahia state: MNHN, Itaparica (praia do Vera Cruz), [1]; MNHN, MD55 sta DC73 (18°59'S / 37°48’W, 607-620 m), Abrolhos Archipel Continental slope, Bouchet, Leal, Metivier coll. [2], — Espirito Santo state: IBUFRJ 8490, off Piama 1993, [3]; MNRJ 10821, off Piuma 1993 [1]; —Rio de Janeiro state: IBUFRJ 6975, GEOMAR XII sta 111, 29/viii/ 1979, NOAC coll., [1]; IBUFRJ 7071, Cabo Frio VII sta 6194 (24° 03,6’ S / 044° 07,6’ W, 134 m), 11/1983, NOAS coll., [1]; IBUFRJ 7836, GEOMAR XII sta 96 (22°05’S / 40°17,4’'W), NOAC coll., [1]. Distribution: USA: Florida (Wise 1996), Texas (Odé 1993); Caribbean: West Indies (Jong & Coomans 1988), Bermuda (Verril & Bush 1900), Bahamas (Redfern 2001); Brazil: Pernambuco state (Mello 1990; this study), Bahia (Oliveira 1992), Espirito Santo state (this study), Rio de Janeiro state (Rios 1994; this study). Remarks: Pseudoscilla babylonia (Figure 3D—E) has been reported from Brazil and other regions in the western Atlantic as Cingulina babylonia (Warmke & Abbott 1962; Rios 1985, 1994; Abbott 1974; Vokes & Vokes 1983; Jong & Coomans 1988; Mello 1990) or Odostomia babylonia (Wise 1996; Redfern 2001). Odé (1993) stated that the genus Cingulina is normally and wrongly used for species from the western Atlantic, and A. D. Pimenta et al., 2007 Page 179 \pleiaalbd eo a Figure 4. A—D. Triptychus niveus Moérch, 1875 (IBUFRJ 14080): A, whole shell (length: 2.7 mm); B—C. protoconchs; D. last whorl; E-I. Triptychus litosbathron n.sp.: E, holotype (MZSP 77065); F—G, paratype (MZSP 77066); H—I, paratype (IBUFRJ 12878); E-F, whole shells (respective lengths: 2.4 mm; 1.6 mm); G—H, protoconchs; I, last whorl. Scale bars: 200 um. introduced the combination Pseudoscilla babylonia, establishing that Pseudoscilla is not related to Cingu- lina. According to Odé (1993), Pseudoscilla is charac- terized by small, regularly conical shells, with sculpture consisting of strongly developed spiral cords, some- times dissolved into separate knobs; Cingulina, on the other hand has more elongate shells, less strongly ornamented, and is restricted to the Pacific Ocean (Odé 1993). Aartsen et al. (1998) reported Pseudoscilla babylonia from the North Atlantic Ocean. However, Penas & Rolan (1999) reviewed the genus Pseudoscilla from West Africa, providing illustrations of the type-species (Pseudoscilla miocaenica), and concluded that the records of P. babylonia by Aartsen et al. (1998) were based on misidentifications of Pseudoscilla bilirata (Folin, 1870). According to Penas & Rolan (1999), Pseudoscilla babylonia is restricted to the Western Atlantic. Subfamily Pyramidellinae Gray, 1840 Genus Triptychus Morch, 1875 Triptychus Morch, 1875: 158. Type species: Triptychus niveus Morch, 1875, by monotypy, St. Thomas. Triptychus niveus Morch, 1875 (Figure 4A—D) Obeliscus (Triptychus) niveus Moérch (1875: 159). Triptychus niveus: Abbott (1974: 300, fig. 3653); Warmke & Abbott (1962: 147, pl. 28e); Vokes & Vokes (1983: 32, pl. 30, fig. 17); Jong & Coomans (1988: 120, pl. 19, fig. 636); Diaz & Puyana (1994: 236, pl. LXIX, fig. 941); Redfern (2001: 145, pl. 65, fig. 603). Triptychus niveus: Rehder (1943: 195, not illustrated) Pyramidella vincta Dall, 1884: Synonimyzed by Abbott (1974). Page 180 Type locality: St. Thomas, Vieques, St. Martin. Material examined: West Indies: IBUFRJ 14080, off Bonaire (12 m), 301.1998, [2]; IBUFRJ 14083, off Bonaire (4m), 14.11.1998, [1]; IBUFRJ 14084, off Bonaire (10 m), 30.1.1998, [1]; —Brazil: -Rio de Janeiro state: IBUFRJ 14078, Arquipélago de Santana, Macaé, v.1983, AG coll., [1]; IBUFRJ 14079, Bacia de Campos sta 22, [1]. Distribution: USA: Florida to West Indies (Abbott 1974; Warmke & Abbott 1962); Mexico: Yucatan Peninsula (Vokes & Vokes 1983); Caribbean: St. Thomas (Morch 1875), Bahamas (Redfern 2001), West Indies (Rehder 1943; Jong & Coomans 1988), Colom- bia (Diaz & Puyana 1994); Brazil: Rio de Janeiro (this study). Triptychus litosbathron n.sp. (Figure 4E—I) Type material: Holotype MZSP 77065, off Parana state, PADCT 6577 (25°15.76'S / 45°04.62'W, 124 m). Paratypes: —Espirito Santo state: IBUFRJ 14081, off Espirito Santo state, REVIZEE sta vv24 (20°S / 34°54"W, 45 m), 27.11.1996, NOAN coll., [1]; MNHN, off Espirito Santo state, REVIZEE sta vv24 (20°S / 34°54'W, 45 m), 27.11.1996, NOAN coll. [1]; IBUFRJ 12878, off Espirito Santo state, REVIZEE sta D1 (20°48'S / 41°09'33"W, 69 m), 23.11.1996, NOAN coll., [1]; MNRJ 10694, off Espirito Santo state, REVIZEE sta D1 (20°48’S / 41°09'33"W, 69m), 23.11.1996, NOAN coll., [1]; —Sao Paulo state: MZSP 86686, REVIZEE sta 6677 (24°40.75'S / 44°50.82'W, 137 m), [1]; MZSP 86688, REVIZEE 6662 (24°00.95'S / 43°55.54'W, 135m), [5]; MZSP 86691, REVIZEE 6666 (24°17.13’S, 44°12.15'W, 163 m), [1]); MNHN, PADCT 6573 (24°42.608’S / 44°43.419’'W, 155 m), [1]; —Parana state: MZSP 77066, type locality, [1]; MNRJ 10941, type locality, [5]; -—Santa Catarina state: MZSP 86693, PADCT sta 6641 (26°15'S / 45°53’W, 130 m) [1]; MZSP 86694, REVIZEE 6695 (26°17.134'S, 46°41.788'W, 153 m), [1]; MZSP 86695, PADCT 6606 (27°48.07'S / 47°24.04’W, 175 m), [1]. Type locality: 25°15.76'S / 45°04.62’W, 124 m; off Parana state, Brazil. Distribution: Only known from Brazil southeast-south coast: Espirito Santo state, Sao Paulo state, Parana state. Etymology: From the Greek /itos: plain, simple; -bathron: base, pedestal. In allusion to the simple, unornamented base of this species. Diagnosis: Small Triptychus species with smooth base, immersed protoconch and small chink-like umbilicus. The Veliger, Vol. 50, No. 3 Description: Shell conic, holotype 2.4 mm in length. Teleconch with up to 5.5 whorls with sinuous outline, due to projections of whorl ornamentation. Suture deep. Protoconch heterostrophic with about 1.5 smooth whorls, with nucleus immersed in first tele- oconch whorl, forming an angle of about 180° with teleoconch main axis; diameter about 250 um. Two deep and wide, channeled spiral furrows, bearing microscopic axial growth lines, one just above the suture, the other in the midline of each teleoconch whorl; between the two furrows, there is a strong, wide, smooth spiral cord; another, stronger spiral cord, just below the suture, is formed by a spiral row of about 18 rounded nodules, axially elongated. Base slightly concave, with microscopic axial ribs and with its adapical periphery marked by a peripheral smooth spiral cord; with a very small chink-like umbilicus, sometimes partially covered. Aperture rhomboid, with a columellar fold medium to weakly projected. Outer lip thin and nearly straight. Remarks: Triptychus niveus has a wide geographic range in the western Atlantic, including localities in the U.S.A. and Caribbean (see distribution for references) and the records of this paper from Brazil (Figure 4A— D), where it has been collected on the southeast coast (about 23°S); Triptychus litosbathron, on the other hand, is restricted to localities on the southeastern and south coasts of Brazil (about 20°—25° S). The inclusion of Triptychus litosbathron (Figure 4E— I) in this genus is somewhat doubtful. The shell has the same general shape, a somewhat more immersed protoconch and similar sculpture, with two spiral cords on each teleoconch whorl, the abapical one is nodulose, the adapical smooth (Figure 4E-F, I); in T. niveus, there are one smooth adapical spiral cord and two nodulose abapical spiral cords on each teleoconch whorl (Figure 4A, D); however, this pattern is not present on the first three adult whorls, which have only one nodulose spiral and the smooth one (Figure 4A— B). In addition, T. litosbathron has an almost smooth base (Figure 41), lacking the spiral cords present in T. niveus (Figure 4D). In spite of all these differences, this seems to be the best generic allocation for the new species herein described, considering also, that it is not desirable to introduce a new generic name in the already confused Pyramidellidae. Diaz & Puyana (1994: pl. LXIX, fig. 936) illustrated a shell named Odostomia (Miralda) sp. The shell is very similar to Triptychus litosbathron, but has an additional spiral cord on the base. Genus Peristichia Dall, 1889 Peristichia Dall, 1889: 339. Type species: Peristichia toreta Dall, 1889, Florida Keys (USA), by original designation. A. D. Pimenta et al., 2007 Page 181 Figure 5. A-—C. Peristichia agria Dall, 1889. A, MNHN; B, MNRJ 10824; C, IBUFRJ 7774; A—B, whole shells (respective lengths: A, 4.0 mm; B, 4.1 mm); C, protoconch; D-H, Peristichia lepta n.sp.: D-G, holotype (MZSP 77062); H, aperture (MZSP 77063). D— E, whole shell (length: 4.2 mm); F, last whorl; G—H, protoconchs. Scale bars: 200 um. Peristichia agria Dall, 1889 (Figure SA—C) Peristichia agria Dall, 1889: 340. Rehder (1943: 195, pl. 20, fig. 4); Abbott (1974: 300. fig. 3655); Diaz & Puyana (1994: 236, pl. LXIX, fig. 942); Vokes & Vokes (1983: 32, pl. 30, fig. 18); Rios (1985: 165, pl. 54, fig. 786, 1994: 188, pl. 62, fig. 879); Mello e Perrier (1986: 139, not illustrated; Mello (1990: 41, fig. 11); Barros (1994a: 74, not illustrated). Type locality: off Cape Hatteras, 63 fms. Material examined: Bahia state: MNHN, Paulista (praia da Concei¢ao), [2]; MNRJ 10824, Itaparica (praia do Despacho), [3]; MNHN, environs de Recife, [2]; MNHN, Cabo (praia de Gaibu), [2]}; MNHN, Itamaraca (praia de Jaguaribe), [4]; MNHN, Cabo (pedras pretas), [3]; MNHN, Paulista (Maria Farinha), [6]; MNHN, Sao Luiz (areia preta), [1]; MNHN, Paulista (Maria Farinha), [1]; MNHN, Cabo (praia de Gaibu), [1]; —Rio de Janeiro state: Col.Mol.UERJ 3338, Ilha Grande sta 15 (Ponta Grande Timuiba, 23°3.762'S 44°36.038’W, 7m), [5]; Col.Mol.UERJ 3337, Ilha Grande sta 16 (Rochedo Sao Pedro, 23°2.868'S 44°32.722'W, 10 m), [1]; IBUFRJ 13686, Angra dos Reis (praia da Figueira), 1998, [1]; IBUFRJ 7774, GEOMAR XII sta 46 (21°30'S, 40°54,8’W, 27 m), vili.1979, [1]; IBUFRJ 2559, Prainha, Arraial do Cabo, [3]. Distribution: USA: North Carolina to Florida (Dall 1889; Abbott 1974); Mexico: Yucatan Peninsula (Vokes & Vokes 1983); Caribbean: Colombia (Diaz & Puyana 1994); West Indies (Rehder 1943); Brazil: Page 182 Ceara to Santa Catarina (Mello 1990; Rios 1994: Barros 1994a). Peristichia lepta n.sp. (Figure 5D—H) Peristichia agria: Farinati (1993: 306, fig. 16). Type material: Holotype MZSP 77062. Paratypes: MZSP 77063, off Sao Paulo state, PADCT sta. 6579 (24°42.302'S, 45°18.831'W, 84 m), [16]; MZSP 77064, off Sao Paulo state, REVIZEE sta 6669 (24°7.42’S, 44°42.22'W, 101 m), [2]; IBUFRJ 9015, Camburi, Espirito Santo state, Eq. Zoo Coll., 15.xii.1987, [2]; IBUFRJ 14085, GEOMAR XII sta 108, 29.viii.1979 NOAS coll., [1]; MNRJ 10693, Ilha Grande sta 36 (Ponta Alta de Parnaioca, 23°12.25’S, 44°30.35'W, 35 m), Rio de Janeiro state, [2]; Col.Mol.UERJ 3335, Ilha Grande sta 36 (Ponta Alta de Parnaioca, 23°12.25’S, 44°30.35’W, 35 m), Rio de Janeiro state, [3]; MNHN, Ilha Grande sta 36 (Ponta Alta de Parnaioca, 23°12.25’S, 44°30.35’'W, 35m), Rio de Janeiro state, [2]. Type locality: off Parana state, Brazil; REVIZEE sta 6656 (25°22.1'S, 46°47.5'W, 70 m). Distribution: Only known from southeast-south coasts of Brazil: Espirito Santo, Rio de Janeiro, Sao Paulo and Parana states; and an additional record from Baia Blanca, Argentina (Farinati 1993). Etymology: From the Latin /eptos: fine, small, thin, delicate. An allusion to the slender shell of this species. Diagnosis: Peristhichia species with slender shell of almost straight whorled outline and base with four smooth spiral cords of equal strength. Description: Shell white, elongate, slender and conical, with about six whorls of almost flat-sided outline; holotype 4.2 mm in length; imperforate. Suture some- what deep. Protoconch heterostrophic helicoid, with about two whorls forming an angle of about 110° to the shell main axis, diameter about 325 um. Axial ribs orthocline, or slightly opisthocline, slender, with upper summits forming small nodules projecting slightly over adapical suture; 22 on body whorl of holotype; interspaces as wide as the ribs, bearing microscopic axial growth lines. Spiral sculpture formed by three narrow cords crossing the axial ribs, forming small rising nodules, well visible on the outline of the whorls, giving rise to a cancellate pattern; median spiral cord stronger than the other two; lower cord forming a channeled furrow below it and above suture. Base somewhat elongate, with about four or five smooth, equally strong spiral cords. Aperture somewhat pyri- The Veliger, Vol. 50, No. 3 form, pointed adapically. Outer lip thin. Columellar fold absent. Remarks: Reheder (1943) discussed the affinities of Peristichia, considering it as a full genus, close to Triptychus. Peristichia lepta (Figure 5D—H) has some similarities to other species from the western Atlantic included in Peristichia, such as Peristichia agria (Figure SA—C) and the type-species Peristichia toreta Dall, 1889 (illustration of specimen by Perry & Schwengel 1955: pl. 23, fig. 160), mainly in the general elongate shell shape, sculpture pattern and protoconch; but differs in the absence of a columellar fold and, mainly, by the approximately four spiral cords on the base, whereas the other species of Peristichia have only one. The most similar species to Peristichia lepta is Peristichia agria, which it resembles in the sculpture pattern, formed by thin, almost orthocline axial ribs crossed by thin spiral cords, giving rise to squared interspaces and small rounded nodules (Figure SA—B). In Peristichia lepta, however, the sculpture is consis- tently weaker, and the shell is more cylindrical and the whorls less convex in outline (Figure 5D—F); whereas in P. agria, the last whorl is somewhat globose, with strongly convex outlines (Figure 5A). Also, in Peristi- chia lepta, there are more numerous and weaker spiral cords on the base (Figure 5F); whereas P. agria has a single, stronger spiral cord in the middle of the base (Figure 5A—B). Peristichia agria has a wide distribution in the western Atlantic, from U.S.A. to southern Brazil (see references in distribution). The known distribution of Peristichia lepta is southeast-south coasts of Brazil. Farinati (1993, 1993: 306, fig. 16) illustrated a specimen of Peristichia lepta, from the Holocene of Bahia Blanca, Argentina, with the name P. agria. This record from Argentina enlarges the geographical and geolog- ical distribution of this new species. Acknowledgments. We are grateful to Dr. Philippe Bouchet and Philippe Maestrati (MNHN) for loan of material from Pernambuco, Brazil; Dra. C. Myaji (Instituo Oceanografico, Universidade de Sao Paulo) for providing additional material from Brazil; Dr. Gary Rosenberg and Mr. Paul Callomon (ANSP) for loan of types, Dr. Robert Hershler (USNM) for sending photographs of type material; Mr. Colin Redfern for comments on the identification of the species and loan of specimens from Bahamas Islands. Dr. Serge Gofas, for review of the manuscript. Dra. Janet Reid, for revising the english version. Mr. J. de Brito (UERJ) and Mr. R. Martins (Cento de Pesquisas da Petrobras SA) for their help with SEM photos. This work was partially supported by ““Conselho Nacional de Desenvolvimento Cientifico e Tecnoldgico” (CNPq). REFERENCES AARTSEN, J. J. VAN. 1984. The pyramidellid-genera described by The Marquis L. de Folin. Bollettino Malacologico 20(5—8):131-138. A. D. Pimenta et al., 2007 AARTSEN, J. J. VAN. 1994. European Pyramidellidae: IV. The genera Eulimella, Anisocycla, Syrnola, Cingulina, Oscilla and Careliopsis. Bollettino Malacologico 30:85—110. AARTSEN, J. J. VAN, E. GITTENBERGER & J. GOUD. 1998. Pyramidellidae (Mollusca, Gastropoda, Heterobranchia) collected during the Dutch CANCAP and MAURITA- NIA expeditions in the South-eastern part of North Atlantic Ocean (part 1). Zoologische Verhandelingen 321: 3-57. ABBOTT, R. T. 1974. American Seashells 2nd ed. Van Nostrand Reinhold Co.: New York. 663 pp, 24 pls. ABSALAO, R. S., F. N. SANTOS & D. DE. O. TENORIO. 2003. Five new species of Turbonilla Risso, 1826 (Gastropoda, Heterobranchia, Pyramidellidae) found off the northeast coast of Brazil (02°—13° S). Zootaxa 235:1—11. ADAMS, A. 1861. On a new genus and some new species of Pyramidellidae from the north of China. Annals and Magazine of Natural History 3(7):295-299. ADAMS, C. B. 1845. Specierum novarum conchyliorum, in Jamaica repertorum, synopsis. Proceedings of the Boston Society of Natural History 2:1—17. ALTENA, C. O. R. 1975. The marine Mollusca of Suriname (Dutch Guiana) Holocene and Recent part III. Gastro- poda and Cephalopoda. Zoologische Verhandelingen 139: 1-104, 11 pls. BARROS, J. C. N. 1994a. Estudo dos componentes bidticos da margem continental brasileira. I Micromoluscos dragados durante a comissao “Canopus,” entre 1965 e 1966. Boletim do Museu de Malacologia 2:57—84. BARROS, J. C. N. 1994b. Moluscos recentes dos recifes costeiros e de sedimentos moveis intertidais de Pernam- buco e da Bahia, Brasil. Cadernos Omega da Universi- dade Federal Rural de Pernambuco (série biologia) 4:35— We BusH, K. J. 1899. Descriptions of new species of Turbonilla of the Western Atlantic fauna, with notes on those previously known. Proceedings of The Academy of Natural Sciences of Philadelphia 51:145—177, pl. 8. CARPENTER, P. P. 1857. Catalogue of the collection of the Mazatlan shells in the British Museum: collected by Frederick Reigen, described by Philip P. Carpenter. Oberlin press, 552 pp. CLENCH, W. J. & R. D. R. TURNER. 1950. The Western Atlantic marine mollusks described by C. B. Adams. Occasional Papers on Mollusks 1(15):233-403. DALL, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Carribean Sea (1879-80), by the U.S. Coast Survey steamer “Blake,” Lieutenant- Commander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. XXIX. Report on the Mollusca. Part 2 Gastropoda and Scaphopoda. Bulletin of the Museum of Comparative Zoology 18:1—492, pls. 10-40. DALL, W. H. & P. BARTSCH. 1904. Synopsis of the genera, subgenera and sections of the family Pyramidellidae. Proceedings of the Biological Society of Washington 17:1— 16. DALL, W. H. & P. BARTSCH. 1906. Notes on Japanese, Indopacific, and American Pyramidellidae. Proceedings of the United States National Museum 30(1452):321—369, pls. 17-26. DALL, W. H. & P. BARTSCH. 1909. A monograph of West American Pyramidellidae mollusks. Bulletin of the United States National Museum 68:xii + 258, 30 pls. Page 183 DIAZ, J. M. M. & M. H. PUYANA. 1994. Moluscos del Caribe Colombiano. Colciencias y Fundacion Natura: Santafeé de Bogota. 291 pp, 37 pls. FABER, M. J. 1988. Studies on West Indian marine molluscs 13. The malacological taxa of Gordon W. Nowell- Usticke. Der Kreukel 24:67—102. FARINATI, E. A. 1993. Pyramidellidae (Mollusca, Gastro- poda) en sedimentos Holocenos de Bahia _ Blanca, Argentina. Ameghiniana 30(3):297-310. FOLIN, L. DE. 1873. En rade de la Pointe-a-Pitre (Guade- loupe). In: L. de Folin & L. Périer (eds.), Les Fonds de la Mer 2:169-171, pl. 6. JOHNSON, R. I. 1989. Molluscan taxa of Addison Emery Verrill and Katharine Jeannette Bush, including those introduced by Sanderson Smith and Alpheus Hyatt Verrill. Occasional Papers on Mollusks 5(67):1—143. JONG, K. M. DE & H. E. COOMANS. 1988. Marine Gastropods from Curacao, Aruba and Bonaire. E. J. Brill: Leiden. 261 pp, 47 pls. LINDEN, J. VAN DER & J. C. A. EIKENBOOM. 1992. On the taxonomy of the Recent species of the genus Chrysallida Carpenter from Europe, the Canary Islands and the Azores (Gastropoda, Pyramidellidae). Basteria 56(1—3):3— 63. MELLO, R. L. S. 1990. Gastropoda: Opistobranchia: Pyrami- dellidae Gray, 1840 da América do Sul, litoral nordeste do Brasil. Caatinga 7:38—43. MELLO, R. L. S. & L. L. PERRIER. 1986. Polyplacophora e Gastropoda do litoral sul de Pernambuco, Brasil. 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La familia Pyramidellidae Gray, 1840 (Mollusca, Gastropoda) en Africa Occidental. 3. El género Chrysallida s. 1. Wberus Suplemento 4:1—73. PENAS, A. & E. ROLAN. 1999. La familia Pyramidellidae Gray, 1840 (Mollusca, Gastropoda, Heterostropha) en Africa Occidental. 6. El genero Pseudoscilla. Wberus 17(2): 11-26. PENAS, A., J. TEMPLADO & J. L. MARTINEZ. 1996. Contribu- cion al conocimiento de los Pyramidelloidea (Gastropoda: Heterostropha) del Mediterraneo espanol. Iberus 14(1):1— 82. PERRY, L. M. & J. S. SCHWENGEL. 1955. Marine shells of the western coast of Florida. Paleontological Research Institution Ithaca. New York. 318 pp. PitsBRY, H. A. & C. G. AGUAYO. 1933. Marine and freshwater mollusks new to the fauna of Cuba. The Nautilus 46(4):116—123. Page 184 PIMENTA, A. D., R. S. ABSALAO & A. S. ALENCAR. 2000. Odostomella carceralis spec. nov. from Ilha Grande, SE Brazil (Gastropoda: Heterobranchia, Pyramidellidae). Basteria 64:65—70. PIMENTA, A. D. & R. S. ABSALAO. 2001a. Taxonomic revision of the species of Turbonilla Risso, 1826 (Gastropoda, Heterobranchia, Pyramidellidae) with type localities in Brazil, and description of a new species. Basteria 65:69— 88. PIMENTA, A. D. & R. S. ABSALAO. 2001b. The genera Bacteridium Thiele, 1929 and Careliopsis Morch, 1875 (Gastropoda: Pyramidellidae) from the east coast of South America. Bollettino Malacologico 37(1—4):41-48. PIMENTA, A. D. & R. S. ABSALAO. 2002. On the taxonomy of Turbonilla puncta (C. B. Adams, 1850) (Gastropoda, Pyramidellidae), with the description of a new species from Brazil and remarks on other western Atlantic species. Zootaxa 78:1—16. PIMENTA, A. D. & R. S. ABSALAO. 2004a. Fifteen new species and ten new records of Turbonilla Risso, 1826 (Gastro- poda, Heterobranchia, Pyramidellidae) from _ Brazil. Bollettino Malacologico 39(5—8):113—140. PIMENTA, A. D. & R. S. ABSALAO. 2004b. 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Spermatophores of six eastern north American Pyramidellid Gastropods and their systematic significance (with the new genus Boonea). Biological The Veliger, Vol. 50, No. 3 Bulletin of The Marine Biological Laboratory 155:360— 382. SA, M. R., J. H. N. LEAL & A. C. S. COELHO. 1984. Gastropodes encontrados no contetdo digestivo de exemplares de Holothuria grisea Selenka, 1867 (Echino- dermata, Holothuroidea) capturados no litoral sul do Estado do Rio de Janeiro, Brasil. Boletim do Museu Nacional 306:3—-12. SCHANDER, C. 1994. Twenty-eight new species of Pyramidel- lidae (Gastropoda, Heterobranchia) from west Africa. Notiziario CISMA 15:11-78. SCHANDER, C., J. J. VAN AARTSEN & J. CORGAN. 1999. Families and genera of the Pyramidelloidea (Mollusca: Gastropoda). Bollettino Malacologico 34(9—12):145—166. TRYON, G. W. 1886. Manual of Conchology; Structural and Systematic, with illustrations of the species. Published by the author: Philadelphia. 8:461 pp., 79 pls. TURGEON, D. D., J. F. QUINN, A. E. BOGAN, E. V. COAN, F. G. HOCHBERG, W. G. LYONS, P. M. MIKKELSEN, R. J. NEvES, C. F. E. ROPER, G. ROSENBERG, B. ROTH, A. SCHELTEMA, F. G. THOMPSON, M. VECCHIONE & J. D. WILLIAMS. 1998. Common and scientific names of aquatic invertebrates from the United States and Canada: mollusks, 2nd edition. American Fisheries, Special Publication 26:526. VERRILL, A. E. 1884. Second catalogue of Mollusca recently added to tha fauna of New England Coast and the adjacent parts of the Atlantic, consisting mostly of deep- sea species with note on other previously recorded. Transactions of the Connecticut Academy of Sciences 6(1):139-294, pls. 28-32. VERRILL, A. E. & K. J. BUSH. 1900. Additions to the marine mollusca of the Bermuda. Transactions of the Connecti- cut Academy of Sciences 10:513—544, pls. 63-65. VOKES, H. E. & E. H. VOKES. 1983. Distribution of Shallow- Water Marine Mollusca, Yucatan Peninsula, Mexico. Middle American Research Institute 54:183, 50 pls. WARMKE, G. & R. T. ABBOTT. 1968. Caribbean Seashells. Livingstone Publ. Co.: Narbeth. xx + 348 pp., 44 pls. WISE, J. B. 1996. Morphology and phylogenetic relationships of certain pyramidellid taxa (Heterobranchia). Malacolo- gia 37(2):443-S51. WIsE, J. B. 2001. Anatomy of Boonea jadisi (Olsson and McGinty, 1958) (Heterobranchia: Pyramidellidae) from the western Atlantic, with comparisons to other species in the genus. The Nautilus 115(2):68—75. THE VELIGER 9) The Veliger 50(3):185-189 (October 1, 2008) © CMS, Inc., 2007 Galba truncatula Miller, 1774 (Pulmonata: Lymnaeidae) in Argentina: Presence and Natural Infection by Fasciola hepatica (Linnaeus, 1758) (Trematoda: Digenea) LAURA ISSIA,' SILVIA M. PIETROKOVSKY,' FLORENCIA KLEIMAN,' PABLO CARMANCHAHP AND CRISTINA WISNIVESKY-COLLI’ "Unidad de Ecologia de Reservorios y Vectores de Parasitos, Departamento de Ecologia, Genética y Evolucion, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Ciudad Universitaria, Pab. H, Cl428EHA, Buenos Aires, Argentina (e-mail: Laura Issia: issia@ege.fcen.uba.ar) ? Asentamiento Universitario de San Martin de los Andes, Universidad Nacional del Comahue, Neuqueén, Argentina Abstract. We report the finding of Galba truncatula in Argentina. In June and August 2006, 157 snails were collected from a stream in Sierras del Palauco, Province of Mendoza, northern Patagonia, Argentina. Fasciola hepatica infection was detected in one of 50 specimens collected in June. In the Americas, this snail was reported from Bolivia and Chile, and its occurrence in Argentina may reflect an ongoing process of geographic expansion. In Argentine Patagonia, Lymnaea viatrix has been regarded as the only lymnaeid involved in the transmission of F hepatica, but our results suggest that G. truncatula may also be playing a role. INTRODUCTION Mollusks are used as first intermediate hosts by all species of digenetic trematodes (Esch et al., 2002). In particular, the cosmopolitan snails of the family Lymnaeidae are involved in the transmission of some digeneans of veterinary and medical importance, such as Fasciola hepatica (Linnaeus, 1758), the causative agent of fasciolosis (Malek, 1985). The endemicity of fasciolosis in a region depends on the presence of intermediate hosts adapted to particular environmental conditions, and therefore the identifica- tion of local lymnaeid species is essential for the design of effective control strategies. In the native fauna of Argentina, species of Lymnaeidae so far reported are Lymnaea diaphana King, 1830; Lymnaea _ pictonica Rochebrune & Mabille, 1885; Lymnaea viatrix Or- bigny, 1835 (Hubendick, 1951; Paraense, 1976, 1982; Malek, 1985; Kleiman et al., 2004); and Pseudosuccinea columella Say, 1817 (Paraense, 1982; Prepelitchi et al., 2003). At present, the only species incriminated in the transmission of F. hepatica are P. columella (Prepelitchi et al., 2003) and L. viatrix, which shows the broadest distribution (Rubel et al., 2005; Cucher et al., 2006; Kleiman et al., 2007). In this work, we report Galba truncatula Miller, 1774 in Argentina, and provide evidence of natural infection of this snail with F. hepatica. MATERIAL AND METHODS The study was performed in a stream at an altitude of 1943 m, located in Sierras del Palauco (35°57'S 69°24’W), Department of Malargiie, Province of Mendoza, northern Patagonia, Argentina (Figure 1). The climate in the study area is cold and arid, with a mean annual temperature of 21.3°C (mean temperature of the coldest month, July: —2.3°C; mean temperature of the warmest month, January: 27.9°C). Winter is the rainy season, and the mean annual precipitation is 198 mm. Biogeographically, the region is included within the Andean—Patagonian domain, Patagonian province, Payunia district, which is characterized by xerophytic vegetation (Cabrera & Willink, 1980). The human population is small and houses are scattered over a large area (density: ). RESULTS Water column densities (number of individuals beneath 100 m* of ocean surface) were computed for each species during day and night periods (Table 1). The combined nighttime density for all species (4388 ind. 100 m-*) was nearly two and one-half times greater than the total daytime density (1777 ind. 100 m~’). Three species (Limacina inflata, Styliola subula, and L. trochiformis) had densities of 1515 to 734 ind. 100 m ~?, Page 195 Wl yy 90 Z Li Ys, 116 _ ip ai ae 5 a a (n= 115) a & se 8 1. ec 80 vo 2 100 x 50 2 (2=7) 140-200 m 50 100 100 50 (n=1) 590-300 m (n= 1) 50 100 ce Taal B 0.6 0.1 mm Size Classes Limacina trochiformis. Legend as for Figure 4. and together represented 68% of the total nighttime density. Eight species (Limacina bulimoides, Creseis acicula, Clio pyramidata, Cavolinia globulosa (Gray, 1850), Limacina lesueuri, Creseis sp., Cavolinia inflexa (Lesueur, 1813), and Creseis virgula (Rang, 1828) conica Eschscholtz, 1829) had nighttime densities between 409 and 45 ind. 100 m °, comprising 30% of the total. The last eight species had nighttime densities of less than 19 ind. 100 m *, making up the remaining 2% of the total. The water column densities during day and night periods were compared by x? analysis (Table 1). Nine of the eleven species with the highest total mean nighttime densities (>45 ind. 100 m *) showed signif- icant diel differences. Only one of those that were significantly different, Limacina trochiformis, was more abundant during the day. A total of five species showed no significant diel differences, and five species were collected only at night. Based on the above results, each species was placed Page 196 The Veliger, Vol. 50, No. 3 Density (ind. 1000 m”) 200 300 Day Night 0-45 m 45-90 m W (n= 1) 140-200 m Percent of Sample oy So Qari nn fn rd [ee ee ee [ reer oe eae oa | er ee a ee | | ee ee Oe Of S | | ll ll | I ll i) SS \ \ NY eLe ie o No} o 5 3 200-300 m 0.05 mm Size Classes Figure 8. Diacria danae. Legend as for Figure 4. into one of two vertical daytime groupings (Table 1, last column). The first group (epipelagic) was com- posed of those species present in comparable densities during both day and night periods with the exception of one, Limacina trochiformis, which was captured in significantly greater numbers during the day. The second group (epipelagic/mesopelagic and mesopelagic) is inferred from the results and, therefore, is hypothet- ical because samples were not collected from the mesopelagic zone. This group consisted of species that were either present in the epipelagic zone in signifi- cantly higher numbers during the night or were absent from the daytime samples. Individual depth interval means and tests for significance for each species in the two groups are summarized in Nigro (2002). Epipelagic Species Group Three species (Creseis sp., C. virgula conica, and Diacavolinia angulosa’ (Eydoux and_ Souleyet, Ms.)(Gray, 1850)) had water column densities that were comparable between day and night (Table 1) and were restricted to the upper 200 m. For Creseis sp., all but four specimens (in the daytime 140-200 m samples) were collected in the upper 140m (Figure 4A). Individuals in the 0-45 m interval were found only at night, and a significantly greater mean nighttime density was found in the 45-90 m interval. The sum of the mean densities below 90 m was about twice as great during the day than at night, but the difference was not significant. Shell lengths ranged from 1.0 to 3.5mm (Figure 4B), and no significant differences were found between diel periods. Creseis virgula conica was limited to the upper 90 m during the day and night (Figure 5A), except for two individuals captured in the nighttime 90-140 m sam- ples. Replicate variability was high above 90 m during both diel periods. Higher daytime densities were recorded from both the 0-45 and 45-90 m depth intervals, but the differences were not significant. Shell lengths ranged from 1.0 to 6.0 mm (Figure 5B). Mean shell lengths were similar in the 0-45 interval, but were significantly greater at night than during the day in the 45—90 m interval. D. T. Nigro & R. R. Seapy, 2007 Density (ind. 1000 m”) 0 1 Mehecse ae 0 5 s0 a. Page 197 0-45 m oO way 5 45-90 m GH [o) | 3 oO = _ 50) (n= 2) 90-140 m 100 50 (n= 1) 200-300 m aap ee La ee 1.8 Dei 0.1 mm Size Classes B Figure 9. Diacria costata. Legend as for Figure 4. Diacavolinia angulosa was recorded in waters above 200 m during both day and night periods (Figure 6A). Mean densities were low (<2 ind. 1000 m~?) with high replicate variability and no significant differences be- tween diel periods. Shell lengths ranged from 2.0 to 4.5 mm and did not differ significantly between day and night periods, although the largest individuals (>3.5 mm) were taken only from the upper 45 m (Figure 6B). Three species (Limacina trochiformis, Diacria dane van der Spoel, 1968, and D. costata Pfeffer, 1879) had daytime water column densities that were similar to their nighttime densities except for L. trochiformis, which had greater densities during the day (Table 1), and extended to 300m. Limacina trochiformis was most abundant above 90 m, with lower densities in the 45-90 m interval and low to very low densities extending down to 300m (Figure 7A). Replicate variability was high with daytime densities higher in all depth intervals above 200 m, but the diel differences were not significant. Shell diameters ranged narrowly between 0.6 and 1.0 mm (Figure 7B). A significantly larger mean shell diameter was found at night in the 0— 45 m interval, but the small (0.02 mm) difference was probably biologically meaningless. Diacria danae was found from the surface to 300 m during the day and was limited to the upper 90 m at night (Figure 8A). Maximal densities were found above 45 m during both day and night periods. Variability was high among replicates, with no significant differ- ences in density between diel periods. Shells ranged narrowly between 1.25 and 1.75 mm (Figure 8B). The largest individuals (>1.60 mm) were found only in night samples, but diel differences were not significant. Diacria costata ranged from the surface to 300 m during the day and was restricted to the upper 90 m at night (Figure 9A). During the day, the majority of individuals were found in the 45—90 m interval, and in the 0-45 m interval at night. Nighttime densities were significantly higher compared to the day in the 0-45 m depth interval. Although daytime densities in the 45— 90 m interval were about three times higher than at night, the difference was not significant. Shell sizes ranged from 1.8 to 2.7 mm (Figure 9B) and were comparable between diel periods in the 0-45 and 45— 90 m depth intervals. Epipelagic/mesopelagic and Mesopelagic Species Group Eight species (Limacina inflata, Styliola subula, L. bulimoides, Creseis acicula, Clio pyramidata, Cavolinia globulosa, L. lesueuri, and Cavolinia inflexa) had Page 198 Density (ind. 1000 m”) 0 50 100 150 200 0 pa 50 i Y///// a 09 Yaa 100 VW a (0.1) Figure 10. Limacina inflata. daytime water column densities that were significantly lower than nighttime densities in the upper 300 m (Table 1). These species are hypothesized to be daytime occupants of the epipelagic/mesopelagic and mesope- lagic zones. Limacina inflata was captured from surface waters to 300 m during both diel periods (Figure 10A), except that it was absent from 140-200 m during the day. At depths above 200m it was recorded in very low numbers during the day. At night replicate variability was high, and significantly greater numbers were captured above 140m with a maximum in the 0-— 45m depth interval. No significant diel differences were found below 140 m, although the mean daytime density in the 200-300 m interval was 17 times greater than at night. Shell diameters ranged from 0.6 to 1.3 mm, and were not significantly different in the 0— 45 m interval (Figure 10B). Small sample sizes (n = 2) in deeper intervals from either day or night periods prohibited statistical comparisons. The Veliger, Vol. 50, No. 3 0-45 m 200 256 45-90 m Ys 2 Q, 5 nN el ° | 3 a 90-140 m r Bieiieee 140-200 m Lancy | (n= 42) 7/ 504 Z 505 (n= 57) ‘i 200-300 m | (n= 2) 50- | 100 4 { aU T T T ] 0.6 13 B 0.1 mm Size Classes Legend as for Figure 4. Styliola subula was present in low to very low numbers above 200 m during the day, and it was restricted to the upper 200 m at night (Figure 11A). It exhibited high replicate variability at night and increased from low numbers at 140-200m to a maximum in the upper 45 m. Significantly greater densities were found only at night in depth intervals above 140 m. The mean nighttime density at 140— 200 m was higher than during the day, but the difference was not significant. Shell lengths ranged broadly from 1.0 to 9.0 mm. No significant differences in shell sizes (Figure 11B) were found, although individuals larger than 5.0 mm were taken only from night samples. Limacina bulimoides was captured from surface waters to 300 m during the day with density increasing with depth, and it was mainly found in the upper 140 m at night (Figure 12A). Nighttime densities were signif- icantly greater compared to the day in depths above 140 m. Replicate variability was high at night with D. T. Nigro & R. R. Seapy, 2007 Density (ind. 1000 m°) 0 20 40 60 £80 = 100 ea =a (0.2) 50 (0.1) 100 S Ss i 150 (ja) 200 300 Day A Figure 11. densities increasing from a low at 140-200 m to a maximum at 045m. The mean daytime density between 140 and 200 m was higher than at night, but the difference was not significant. Shell lengths ranged narrowly from 0.6 mm to 1.6 mm and were signifi- cantly larger at night in the 90-140 m depth interval (Figure 12B). In waters above 90m, the largest individuals (>1.4 mm) were taken only at night. Creseis acicula was present in low numbers from the surface to 300 m during the day and was restricted to waters above 200m at night, although it was only abundant above 90 m (Figure 13A). High replicate variability was found among the night tows above 90 m. Nevertheless, nighttime densities were signifi- cantly greater than daytime densities in the 0-45 and 45-90 m intervals. Most (80%) of those individuals captured at night were in the 0-45 m interval. Shell lengths ranged broadly from 2.0 to 9.0mm (Fig- ure 13B) with no significant diel differences, except in the 45-90 m interval where the mean length was significantly larger at night. Clio pyramidata was absent from waters above 140 m and was scarce (mean abundances < 1.0 ind. Page 199 0-45 m Y Yi o Gj Ge 655) 504 158 45-90 m ' 1 a. =| 90-140 m nN oo fo} 5 2) Z 30 | (n= 4) 4] 7 vee 140-200 m 301 -_ (n=9) | U7 604 60 200-300 m if T T T ] T [fae | 1.0 9.0 B 1.0 mm Size Classes Styliola subula. Legend as for Figure 4. 1000 m~*) between 140 and 300m during the day (Figure 14A). At night, this species was present only above 200 m with a maximal density in the 90-140 m interval. The total mean nighttime density was 20 times greater than during the day. Shell sizes varied widely from 1.0 to 13.0 mm (Figure 14B). Only the smallest shells (1.0-2.0 mm) were recorded from day tows. Large shells (>7.0 mm) were all recorded from night tows in the upper 90 m. Cavolinia globulosa was present down to 140 m during the day and to 300 m at night (Figure 15A). Mean nighttime densities were significantly greater than daytime densities in the 0-45 m interval and significantly less in the 45-90 m interval. No significant differences were found below 90m. Shell lengths ranged broadly from 0.5 to 6.5mm and _ were significantly greater at night in all depth intervals above 140 m (Figure 15B). Large individuals (>3.5 mm) were collected only at night from tows above 200 m, except for one shell (5.5 mm) collected during the day from the 90-140 m interval. Limacina lesueuri was present in very low numbers (<0.3 ind. 1000 m~?) between the surface and 300 m Page 200 The Veliger, Vol. 50, No. 3 Density (ind. 1000 m®) 0-45 m © 10) 20, 530° 40 50" 0 20 40 50 (0.1) _ _— l//// / 504 50 ee a 45-90 m (0.4) _ An 77 Li l _ oe A 100 7/7 E £ ‘5 407 (n= 41) é 150 : 4 ee 90-140 m : Z hoe ara 200 140-200 m 300 Day Night 40 5 (n= 120) ‘i | Bel L.. 200-300 m Figure 12. during the day and was recorded in increasing numbers from 200m to the 0-45 m interval at night (Fig- ure 16A). Total mean nighttime densities were 52 times greater than daytime densities. Sixty-three percent of those captured at night were from the 0-45 m interval. Shell sizes ranged narrowly from 0.7 to 1.3 mm (Figure 16B) with the largest (>1.2 mm) captured only at night above 140m. Many zero density samples prevented statistical comparison of density and size distributions. Cavolinia inflexa was captured down to 300m during the day and to 200 m at night (Figure 17A). Although slightly higher mean densities were recorded in all depth intervals at night, no significant differences were found, most probably due to the high replicate variability. Shells lengths ranged broadly from 1.0 to 6.0 mm. Those >2.5 mm were found only in the upper 140 m at night (Figure 17B), and individuals from the intervals above 90 m were significantly larger at night. Five species (Cuvierina columnella (Rang, 1827), Hyalocylis striata (Rang, 1828), Diacria maculata Bleeker and van der Spoel, 1988, Cavolinia gibbosa if T ip T T T Tae Tecoma) 1 1.6 B 0.1 mm Size Classes Limacina bulimoides. Legend as for Figure 4. (d’Orbigny, 1836), and D. major (Boas, 1886)) were absent from the upper 300 m during the day (Table 1). These species are hypothesized to be daytime occupants of the mesopelagic zone. Except for Cuvierina colum- nella and Hyalocylis striata, which were captured in moderate to low numbers at night, the remaining three were captured in extremely low numbers at night and were not recorded from day tows. C. columnella was captured in the upper 200 m (Figure 18A), and most individuals (64%) were recorded from the 0-45 m interval. Shell lengths ranged narrowly between 7.0 and 8.0 mm (Figure 18B) with the largest individuals (>7.6 mm) between 45 and 90 m. Hyalocylis striata was recorded only in waters above 90 m at night (Figure 19A). Mean densities were low (<1.0 ind. 1000 m~?) and nearly the same in the 0-45 and 45—90 m intervals. Shell length ranged from 2.5 to 7.0 mm (Figure 19B) with shells larger than 3.5 mm taken only from the 45—90 m interval. Diacria maculata and D. major were captured in extremely low numbers (five and one individuals, respectively) above 140m at night. Sizes were not D. T. Nigro & R. R. Seapy, 2007 Density (ind. 1000 m”) 0 10 20 30 40 #50 0 10 20 30 50 100 150 Depth (m) 200 300 Day Night Figure 13. measured, as all shells were broken to some extent making size estimation unreliable. Lastly, a single Cavolinia gibbosa with a shell length of 9.2 mm was captured at night from the 200-300 m depth interval. DISCUSSION Based on the reviews of euthecosome taxonomy and biogeography by van der Spoel (1967), Bé and Gilmer (1977), and van der Spoel et al. (1997), and two subsequent ‘taxonomic studies (van der Spoel and Pierrot-Bults, 1998 and Bontes and van der Spoel, 1998), 68 species of euthecosomes are currently recognized from the world’s oceans. Half (34) have been reported from the tropical and/or subtropical Pacific Ocean (Table 2). McGowan (1960, 1963, 1971) identified 18 species with distributions that coincide with Hawaiian waters, all of which were collected in the present study with two exceptions. The first, Cavolinia uncinata (Rang, 1829), was classified as a tropical species by Be and Gilmer (1977) and may be limited to lower latitudes south of the Hawaiian Islands. The second, Cavolinia tridentata (Neibuhr, 1775), was classified as subtropical and uncommon by Tesch (1948) and McGowan (1960). Of the 19 species identified from the upper 300 m of the water column in the present study, 8 were present in a UZ Page 201 0-45 m 50 80 45-90 m oO E N Gey fo) =I 8 2 90-140 m 50 Ewen= 200-300 m (esa See Waa Tae | 2.0 9.0 1.0 mm Size Classes B Creseis acicula. Legend as for Figure 4. significantly greater numbers at night than during the day while 5 were collected only at night, and, by implication, were below 300 m during the day. The most probable explanation for the difference in day- night densities for these 13 species 1s nocturnal vertical migration. If a major adaptive value for nocturnal vertical migration is reduced visibility to visual predators during the day, then the shallowest daytime depths should coincide with that depth at which an individual ceases to present a perceptible visual cue (Angel, 1985). Off Hawaii, the shallowest daytime depth at which counterillumination can occur is 400 m, which coin- cides with the shallowest daytime depth of midwater micronekton (Young et al., 1980). This depth also should coincide with the shallowest daytime depth for mesopelagic euthecosomes and could explain the absence or low abundance during the day of the 13 species above 300 m in the present study. An alternate hypothesis explaining differences in density between day and night periods is the ability of animals to avoid an oncoming plankton net during the day. Daytime net avoidance was hypothesized by McGo- wan and Fraundorf (1966) to be a function of net size. They found that 20- and 40-cm diameter nets (0.03 and 0.13 m*) underestimated euthecosome abundances ob- tained with a 140-cm net (1.54 m7). Nets with diameters Density (ind. 1000 m?) 50 100 € a iS 150 QA 200 300 Day A Figure 14. of 60, 80 and 100 cm (0.28, 0.50, and 0.79 m?*) gave intermediate results. Hypothetically, net avoidance of the 70-cm Bongo nets used here should be comparable with that obtained with their 60- and 80-cm nets. However, the paired 70-cm Bongo nets have unob- structed mouth openings while the individual nets used by McGowan and Fraundorf employed three-point towing bridles. Strong support for an hypothesis that diel differences in density of euthecosomes at epipelagic depths are due to vertical migration and not net avoidance 1s given by data from Wormuth (1981). He compared the density of nine abundant euthecosomes from 22 day and night samples collected from nine discrete depth intervals to 1000 m using a l-m? multiple open-closing net and environmental sensing system (MOCNESS). Like the Bongo nets, the mouth opening of the 1-m?* nets was unobstructed. Wormuth found no significant difference between total species densities from day and night periods. Additional support for the above hypothesis is the lack of eyes or well-developed photoreceptors that the; Veliger, Voly50) Noms WU 0-45 m a 45-90 m L, | oe 90-140 m ovo E n GH ° r= 3 By a al 140-200 m 100 50 (n= 8) 200-300 m 1.0 13.0 B 1.0 mm Size Classes Clio pyramidata. Legend as for Figure 4. would be sufficiently sensitive to detect a change in light intensity produced by an oncoming net. Eutheco- somes do possess one (in Limacinids) or two tentacles with tissue that may serve to detect differences in light intensity (Lalli and Gilmer, 1989). However, their ability to detect an oncoming net is improbable. A tactile function of the tentacles is probable, but such sensory receptors would operate equally as well during the day as at night to invoke net avoidance behavior. For some species, it is also possible that larger individuals are more capable of daytime net avoidance than smaller ones. In the present study, we found that daytime tows in the upper 300 m collected only small individuals of Cavolinia inflexa (<2.5 mm) and Clio pyramidata (<2.0 mm), while nighttime tows collected individuals up to 6.0 mm and 13.0 mm, respectively. However, an alternate explanation is that resident daytime populations of small animals reside between the surface and 300 m, while larger individuals are found deeper. Daytime and nighttime patterns of vertical distribu- D. T. Nigro & R. R. Seapy, 2007 Density (ind. 1000 m°) 0 10 20 30 wy 50 100 150 Depth (m) 200 300 Day Figure 15. tion would be expected to differ between localities mainly as a result of differences in light penetration, which is affected by water turbidity and season (at increasingly higher latitudes). Even in some tropical and subtropical areas, water clarity may vary due to seasonal upwelling and river runoff. In the eastern Caribbean, for example, massive freshwater runoff from the Orinoco River can cause dramatic changes in turbidity. This phenomenon has been observed on a seasonal basis far from the river mouth in the waters around Barbados (C. Lalli, personal communication). Secondary causes of variability in the depth of the epipelagic zone include increased turbulence due to storms, variations in light penetration due to atmo- spheric conditions, and variability in plankton density. For example, the samples used in the present study were collected during a period of seven days in the month of April with clear weather and calm seas. It is reasonable to expect that samples taken during other times of the year of the study may have yielded different results from those reported here. Lacking Page 203 50 : 0-45 m GY 50 80 a 42.8 40 H/o 40 80 60 30 90-140 m Percent of Sample Ss KK “Yy Ws, 30 60 — a 200-300 m 1.0 mm Size Classes Cavolinia globulosa. Legend as for Figure 4. information regarding the depth of the epipelagic zone at the localities and times of the previous studies in the North Atlantic (discussed below), the vertical distribu- tion patterns characterized here from Hawaiian waters may or may not be directly comparable. Nonetheless, we decided that we should examine intra-species patterns reported by previous authors whose studies were conducted in the western North Atlantic with the aforementioned aspects serving as an explanatory framework for differences that may occur. Among the six species classified as epipelagic in the present study, three (Limacina trochiformis, Creseis virgula conica, and Diacavolinia angulosa) showed no evidence of migration, while three (Creseis sp., Diacria danae, and D. costata) performed limited nocturnal migrations. Wormuth (1981) found that L. trochiformis and C. virgula conica from the Sargasso Sea were non- migrators with peak abundances above 100 m and 200 m, respectively, during both day and night periods. These results are similar to those of the present study, where >50% of the sampled population of L. Page 204 The Veliger, Vol. 50, No. 3 50 ss yy Uy yy WY rn beers 0-45 m Density (ind. 1000 m®) 20 —__// 0. S 10 158 20) Fost) 0m se IORMEIONNE DOS mmEns( (Se) 100 7/ E € ‘S € 150 5 a & 200 a — 7 140-200 m (0.2) : 50 300 Day Night | i Ge2 200-300 m oe) ee 13 B 0.1 mm Size Classes Figure 16. Limacina lesueuri. Legend as for Figure 4. trochiformis was in the upper 140 m and no individuals of C. virgula conica were found below 140 m during either diel period. Downward nocturnal migration from the surface by adults of C. virgula conica in the Caribbean were reported by Haagensen (1976). There may be some evidence of this pattern in the present study, as nighttime densities were somewhat lower than daytime densities above 90 m. The vertical distribution of ‘“Cavolinia longirostris”’ was reported by Wormelle (1962). She found evidence of migrations in the Florida Current with 50% of the individuals above 219 m during the day and above 76 m at night. In contrast, Chen and Bé (1964) found no diel differences in surface waters (0-10 m) in the western North Atlantic. Comparison of these results with the present is not possible because “C. /ong- irostris’’ was split into 24 species by van der Spoel et al. (1993). One of these species, Diacavolinia angulosa, was identified in the present study. Which species (or multiple species) was represented by “‘C. Jongirostris” in Wormelle’s study is not known. Our results for Diacria danae and D. costata can be compared indirectly with three reports for D. quad- ridentata (de Blainville, 1821), which may actually represent one or both of the above species (discussed in Nigro, 2002). Off Hawaii, D. danae showed a pattern that was nearly identical to D. costata, remaining exclusively in the upper 140 m during both diel periods. There was some evidence of migration by individuals dwelling deeper than 90 m during the day to waters above 90 m at night, which is in general agreement with patterns found in the Caribbean by Haagensen (1976) and in the western North Atlantic by Wormelle (1962). However, in the surface waters of the western North Atlantic, Chen and Bé (1964) found no evidence of migration for D. quadridentata. Ten of the 13 species assigned to the epipelagic/ mesopelagic and mesopelagic species group appeared to undergo nocturnal vertical migrations in the present study; three Limacinidae (Limacina inflata, L. buli- moides, and L. lesueuri), and seven Cavoliniidae (Styliola subula, Creseis acicula, Clio pyramidata, Cavolinia globulosa, Cavolinia inflexa, Cuvierina colum- nella, and Hyalocylis striata). These species were either absent or present in low numbers in the upper 300 m during the day and in moderate to high numbers in the upper 140 m at night. Wormuth (1981) recorded L. inflata down to D. T. Nigro & R. R. Seapy, 2007 Density (ind. 1000 : 0 1 2 3 4 3 a 50 = __ 200 300 Day Night Figure 17. =1000 m in the Sargasso Sea during the day with peak densities between 100 and 400 m. He found most of the individuals at night in the upper 75 m with a gradual decrease in numbers down to 125 m. In the present study, the total daytime density in the upper 300 m was 3.2% of the nighttime density. Assuming that night tows are a reasonable estimate of population density and that daytime net avoidance is either absent or not significant, then only 3.2% of the population occurred above 300 m during the day whereas >50% were found at depths above 100 m at night. These data correspond generally with the results described above from Wormuth from the Sargasso Sea. In marked contrast, Wormelle (1962) reported that in the Florida Current 50% of the individuals were captured above 236 m during the day and 232 m at night (i.e., no evidence of nocturnal migration). Styliola subula and Limacina bulimoides were found by Wormuth (1981) to be vertical migrators in the Sargasso Sea, with most of each species population above 100 m at night and above 260 m during the day. In the present study, the water column density of S. Page 205 0-45 m 100 3-4 6.1 L| | 45-90 m 7.4 ) E 90-140 m n (ray jo) ra 3 o a 140-200 m 200-300 m fee =e T T =I Lm aL La) 1.0 6.0 B 0.5 mm Size Classes Cavolinia inflexa. Legend as for Figure 4. subula was 13 times greater at night than during the day (Table 1). Also, most of the specimens from the day samples were 1—3 mm, while those from the night tows ranged from 1-9 mm (Figure 13). These density and size differences suggest nocturnal migration of S. subula greater than 3 mm from depths below 300 m. For L. bulimoides, there were no diel differences in shell sizes, although in agreement with Wormuth there was an evident nocturnal vertical migration from waters below 140m during the day to above 140m at night. Wormuth (1986) reported that shell size increased with decreasing depth in the upper 100 m; most individuals ranged in size from 0.5 to 0.8 mm between 50 and 100 m, 1.2 to 1.4 mm between 25 to 50 m, and 1.3 to 1.5 mm from 0 to 25 m. Comparison of shell sizes in the 0-45 and 45—90 m intervals in our study, however, show no such vertical difference, especially at night where the size-frequency distributions were based on large sample sizes (Figure 14). Myers (1967) reported maximum concentrations of Creseis acicula at night in the upper 50 m off Cape Hatteras, and Wormuth (1981) suggested that some Page 206 The Veliger, Vol. 50, No. 3 0-45 m Density (ind. 1000 m™®) 2 3 4 5 = rs 50 V/ EB - 3 100 5 5 a Y 140-200 m a ‘_ é 50 200 4 / 100 _ 7.0 8.0 300 0.1 mm Size Classes B A Figure 18. Cuvierina columnella. Legend as for Figure Density (ind. 1000 m®) 0-45 m Percent of Sample Z YY Y Y 0.5 mm Size Classes an Night Figure 19. Hyalocylis striata. Legend as for Figure 4. D. T. Nigro & R. R. Seapy, 2007 Page 207 Table 2 Geographic distribution of euthecosomes and comparison of species collected from the North and Equatorial Pacific Ocean by McGowan (1960) and the present study (indicated by asterisks). Only distinct species are listed; for those with known morphological variants (i.e., subspecies, varieties, or formae), the distribution includes all subspecific taxa. Species in bold print have tropical and/or subtropical distributions in the North Pacific Ocean. Sources of data are: 1) Bé and Gilmer (1977), 2) van der Spoel et al. (1993), 3) van der Spoel et al., (1997), 4) Bontes and van der Spoel (1998), and 5) van der Spoel and Dadon (1999). + = present, — = absent. Biogcostapbie Region Source of Mc Gowan Species Polar Subpolar Subtropical Tropical Data (1960) Family Limacinidae + Ww Nn + Limacina bulimoides* ~ = a Limacina helicina + Limacina helicoides — _ ae Limacina inflata* _ = ae Limacina lesueuri* — = ae Limacina retroversa ~ A = Limacina trochiformis* - _ ae + | | hia + + Family Cavoliniidae Cavolinia gibbosa* = = Cavolinia globulosa* al ae Cavolinia inflexa* = = Cavolinia tridentata = = Cavolinia uncinata — — Clio andrae re = Clio antarctica = + Clio balantium = = Clio campylura = = Clio chaptalii = = Clio convexa = = Clio cuspidata = = Clio polita = + Clio pyramidata* = = Clio recurva = = Clio scheelei = = Clio sulcata = + Creseis acicula* = = Creseis chierchiae = = Creseis sp.* = = Creseis virgula = = Cuvierina columnella* = = Diacavolinia angulosa* = = Diacavolinia elegans — = Diacavolinia longirostris = = Diacavolinia mcgowani = a Diacavolinia pacifica = = Diacavolinia triangulata = = Diacavolinia vanutrechti = = Diacria costata* =; = Diacria danae* = = Diacria maculata* = = Diacria major* = = Diacria quadridentata = = Diacria rampali = = Diacria schmidti = = Diacria trispinosa = = Hyalocylis striata* = = Styliola subula* i = Je Gp ch Yee | eB) ae et Pe (eeeeeeeerace nA n ee fees db Sa) hk aes ;t+tt¢4 CO a tO a Oe 0 NAwn Nn | | ++ | + + Plt) +tet tt Y N | + ++ + | et |} +++ on | | aE oy | +++ ttettere t+ +e tee tees Www Un nn GPa op KH HK wWBWRWARWWNNNNNNNYH | | +++ Page 208 shallow water migration by this species might take place to waters above 25 m at night in the Sargasso Sea. These findings agree with our results; at night more than 95% of the individuals were in the upper 90 m and 70% were in the upper 45 m. In both of the preceding studies, the daytime distribution was con- centrated in the upper 100 m, while we found low daytime densities between the surface and 300 m. Wormuth (1981) recorded Clio pyramidata down to 1000 m with peak abundances between 240 and 460 m during the day and in the upper 100 m at night. We found that only 4% of the individuals captured at night between the surface and 300 m were present in this depth range during the day, which is consistent with Wormuth’s findings from the Sargasso Sea. The results of the two studies differ in that most of the nighttime population in the present study was between 100 and 200 m instead of the upper 100m as Wormuth reported. The presence of small (1.0—1.1 mm) individ- uals during the day between 140 and 300 m, a broad size range (to 13.0 mm) at night, and those greater than 6 mm only collected in the upper 90 m at night strongly suggests nocturnal vertical migrations by the adult members of this species. Limacina lesueuri was reported by Wormuth (1981) to be a nocturnal migrator with 50% of the individuals above 300 m during the day and above 100 m at night. Likewise, Haagensen (1976) reported more than 90% of the individuals above 274m during the day and above 63 m at night. We found that the total daytime density was only 2% of the total nighttime density above 300 m, implying that most individuals in the population were deeper than the nets fished. At night, >50% of the individuals were in the upper 45 m. Thus, the daytime vertical range off Hawaii may be deeper than in the Sargasso Sea and the Caribbean; however, the nighttime depths of greatest abundance are similar. Cavolinia inflexa was found by Haagensen (1976) to be a vertical migrator in the Caribbean, with 50% of the population sampled above 261 m during the day and 50% above 50m at night. Similar results were obtained in this study, as the daytime density above 300 m was 40% of the night density. At night, >50% of the population sampled were found above 90 m. In the Caribbean, Haagensen (1976) reported migra- tions of Cuvierina columnella from a depth range of 224 to 344m during the day to above 65m at night. Similarly, he found 50% of Hyalocylis striata above 243m during the day and above 81m at night. Wormelle (1962) reported similar findings for H. striata in the Florida Current with 50% above 283 m during the day and 84 m at night. In the present study, both species were absent from daytime tows and were limited to the upper 90 m at night, except for one C. columnella captured in the 150-200 m interval. During the day, the Bongo nets may have simply missed The Veliger, Vol. 50, No. 3 individuals that were there or the nets did not fish deep enough to reach the minimum daytime depth of this species. Like the present study, Myers (1967) reported C. columnella and H. striata off Cape Hatteras to be absent during the day from tows above 150 m, but found both species to be concentrated in the upper 50 m at night. Three (Diacria maculata, Cavolinia gibbosa, and Diacria major) of the 19 species of euthecosomes identified from Hawaiian waters were captured only at night in extremely low numbers, and comparisons with other studies are not warranted. CONCLUSIONS The diel distributions of the 16 most abundant species discussed above are in general agreement with the results obtained for the same species that occur in the North Atlantic Ocean and Caribbean Sea. Six of the species sampled here were permanent residents of the epipelagic zone and showed limited to no diel differences. The majority (10) were either absent or present in low numbers in the upper 300 m during the day and in higher numbers in the upper 140 m at night. If these low densities, or absence during the day, are a result of most of the species’ populations residing below 300 m, then these ten species generally dwell in deeper waters off Hawai than in the North Atlantic and Caribbean. Acknowledgments. The support of the officers, crew, and members of the scientific party during cruises of the R/V KANA KEOKT are gratefully acknowledged. The cruises were undertaken in conjunction with R.E. Young of the University of Hawaii, whose collaboration is greatly appreciated. Thanks are also due to L. Newman for help with Creseis identifica- tions, and to K. Messer who provided valuable help with the statistics. LITERATURE CITED ANDERSON, V., J. SARDOU & B. GASSER. 1997. 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Variation in Cavolinia inflexa (Lesueur, 1813) (Gastropoda:Pter- opoda:Euthecosomata). Zoologische Verhandelingen (Leiden) 323:435—440. WORMELLE, R. L. 1962. A survey of the standing crop of plankton of the Florida Current. VI. A study of the distribution of the pteropods of the Florida Current. Bulletin of Marine Science of the Gulf and Caribbean 12: 93-136. WoRMUTH, J. H. 1981. Vertical distributions and diel migrations of Euthecosomata in the northwest Sargasso Sea. Deep-Sea Research 28:1493-1515. WoORMUTH, J. H. 1986. Factors affecting the biogeography of mid to low latitude euthecosomatous pteropods. Pp. 270— 277 in A. C. Pierrot-Bults, S. van der Spoel, B. J. Zahuranec & R. K. Johnson (eds.), Pelagic biogeography: proceedings of an international conference, The Nether- lands 29 May — 5 June 1985. UNESCO technical papers in marine science 49. YOUNG, R. E. & R. F. HARMAN. 1985. Early life history stages of enoploteuthin squids (Cephalopoda: Teuthoi- dea: Enoploteuthidae) from Hawaiian waters. Vie Milieu 35(3/4):181—201. YOUNG, R. E., E. M. KAmpPA, S. D. MAYNARD, F. M. MENCHER & C. F. E. ROPER. 1980. Counterillumination and the upper depth limits of midwater animals. Deep-Sea Research 27:671—691. The Veliger 50(3):210-218 (October 1, 2008) THE VELIGER © CMS, Inc., 2007 Two New Species of Doriopsilla from the Tropical Western Atlantic with Remarks on Cariopsillidae Ortea & Espinosa, 2005 ANGEL VALDES Department of Biological Sciences, California State Polytechnic University, 3801 West Temple Avenue, Pomona, California 91768, USA (e-mail: aavaldes@csupomona.edu) JEFF HAMANN 1000 Pioneer Way, El Cajon, California 92020, USA Abstract. Four Caribbean species of Doriopsilla are described, including two new species to science. The first detailed anatomical examinations and illustration of living Caribbean animals of Doriopsilla areolata and Doriopsilla nigrolineata reveals that these are two distinct species. Two undescribed species are differentiated from previously known taxa by a combination of external and anatomical characteristics, including the structure of the digestive and reproductive systems, the presence or absence of black lines and white spots on the dorsum, and the morphology of the dorsal tubercles. The genus Cariopsilla and the family Cariopsillidae are synonimized with Doriopsilla and Dendrodorididae respectively, based on the application of modern principles of systematic biology. INTRODUCTION Valdés and Ortea (1997) revised the species of the genus Doriopsilla in the Atlantic Ocean, regarding only three species as valid. Doriopsilla areolata Bergh, 1880, with three subspecies, is distributed through Southern Europe (D. areolata areolata), West Africa (D. areolata albolineata) and the Caribbean Sea (D. areolata nigrolineata). Doriopsilla pelseneeri d’Oliveira, 1895 is only present in the Iberian Peninsula, and Doriopsilla pharpa Marcus, 1961, is known from the Atlantic coast of North America and the Caribbean Sea. Since then, a third Caribbean species, Doriopsilla espinosai Valdés & Ortea, 1998 was described based on material collected from Cuba (Valdés & Ortea, 1998). More recently, Ortea & Espinosa (2005) erected the new genus Cariopsilla Ortea & Espinosa, 2005 and the new family Cariopsillidae Ortea & Espinosa, 2005 for Doriopsilla pharpa based on the presence of caryophyl- lidia-looking tubercles in this species. Valdés et al. (2006) illustrated several additional undescribed species from the Caribbean and brought Doriopsilla nigroli- neata back to the species level based on consistent morphological differences with Doriopsilla areolata. The present paper deals with the description of two of the new species illustrated by Valdés et al. (2006) and discusses the status of Cariopsillidae based on modern concepts in systematic biology. All specimens are deposited at the Malacology Section of the Natural History Museum of Los Angeles County (LACM). SPECIES DESCRIPTIONS Doriopsilla elitae Valdés & Hamann, n. sp. (Figures 1A—C, 2, 3A) Doriopsilla sp. 1 — Valdés et al., 2006: 204-205. Material examined: HOLOTYPE: Vieux Fort, South Point, St. Lucia, 30m depth, 1 October 1987, 1 specimen 20 mm long, live (LACM 1930). PARA- TYPE: Petit Nevis, St. Vincent and the Grenadines, 18 m depth, 18 January 1987, 1 specimen 20 mm long, live (LACM 1931). Additional specimens were photo- graphed but are not preserved. External morphology: Living animals reach up to 20 mm in length. The general color of the living animals is variable from yellow to dark orange (Figure 1A—C). The dorsum is covered with opaque white patches, generally small, and situated on top of many dorsal tubercles. Some of the patches are much larger than the rest, also covering large tubercles situated along the edge of the visceral hump. The rhinophores and gill are the same color as the rest of the body. The body is flat, oval (Figure 1A—C), stiffened by a subepidermal network of strong spicules over the entire body surface. The dorsum is covered by a number of low, simple, conical tubercles stiffened with spicules. The mantle margin is wide and slightly undulate. The rhinophores are perfoliate with up to 13 lamellae. The A. Valdés & J. Hamann, 2007 Ch Pa = Ee Figure 1. Living animals. A, Doriopsilla elitae from St. Vincent, 19 mm long (LACM 1931) — photo Jeff Hamann. B, Doriopsilla elitae from Martinique, size unknown — photo Greg Hamann. C, Doriopsilla elitae from Aruba, 25 mm long — photo Jeff Hamann. D, Doriopsilla tishae from Roatan, Honduras, 49 mm long (LACM 1933) — photo Jeff Hamann. E, Doriopsilla areolata from Martinique, 36 mm long (LACM 173780) — photo Jeff Hamann. F, Doriopsilla nigrolineata from Guanaja, Honduras, 25 mm long (LACM 173781) — photo photo Jeff Hamann. The; Veliger; Vol: 50; Noms Figure 2. Drawings of the internal anatomy of the paratype of Doriopsilla elitae (LACM 1931). A, General view of the anatomy, scale bar = | mm. B, Detail of the anterior portion of the digestive system, scale bar = | mm. C, Reproductive system, scale bar = | mm. D, Connection of the bursa copulatrix and seminal receptacle, scale bar = 1 mm. Abbreviations: am, ampulla; bb, buccal bulb; be, bursa copulatrix; bg, blood gland; dd, deferent duct; dg, digestive gland; e, esophagus; fg, female gland; ht, heart; i, intestine; pr, prostate; rm, retractor muscle; rs, reproductive system; s, syrinx; sr, seminal receptacle; v, vagina. gill is composed of four tripinnate leaves. The anus is eccentric to the left. The oral tentacles are fused together with the mouth opening in the center. The anterior border of the foot is slightly concave but not notched. Anatomy: The buccal bulb is oval (Figure 2A—B), covered by minute, rather undifferentiated oral glands on its proximal portion. The tubular esophagus leads from the buccal bulb. The esophagus is very long and convoluted (Figure 2A). Posteriorly, it broadens into a short muscular portion. The intestine runs posteriorly in the usual position and lacks any pyloric gland (Figure 2A). The ampulla is short and muscular (Figure 2C). It divides into a short oviduct, which enters the female gland, and the prostate. The prostate is broad and flattened. From its distal end, the prostate leads into an elongated and convoluted deferent duct. The penis, when everted, is very long and contains several rows of penial hooks. The penial hooks are approximately 40 um wide at the base and up to 55 um in length (Figure 3A). The vagina is long and straight. At its proximal end is a large, thin-walled, oval bursa copulatrix. The seminal receptacle is small, having a long duct that joins the vagina at the point where it connects the bursa copulatrix. From this point also emerges the uterine duct. The circulatory system consists of a large heart (Figure 2A), joined by the aorta with a flattened blood gland, situated behind the central nervous system. Etymology: The species is named after Elita Hamann, middle daughter of Jeff Hamann. Geographic range: Thus far this species has been collected or photographed in Aruba, St. Vincent and the Grenadines, Grenada, and Martinique. Remarks: Doriopsilla elitae is clearly different from other Atlantic species of the genus. There are no other species with yellow or red background colorations and solid white spots. Species with uniform white, yellow, orange or red colorations include Doriopsilla areolata and Doriopsilla pelseneeri d’Oliveira, 1895, but the former has a complex pattern of white rings or lines and the latter lacks white pigment except for a single ring around the gill pocket. Anatomically, Doriopsilla areolata is clearly different by having a longer, thinner ampulla, a wider prostate, a proportionally larger seminal receptacle, and a much more elongate buccal bulb. Doriopsilla pelseneeri lacks any white pigment on the dorsum, except for a white line around the branchial sheath (Valdés & Ortea, 1998). Additionally, D. pelseneeri has large, irregular dorsal warts that contrast with the low, simple, conical tubercles of D. elitae. For a comparison of the external characteristics of D. elitae with other Atlantic species of Doriopsilla see Table 1. Eastern Pacific species of Doriopsilla with yellowish to reddish background color and white spots include Doriopsilla albopunctata (Cooper, 1863) and Doriopsilla gemela Gosliner, Schaefer, & Miller, 1999), both characterized by having very small opaque white spots uniformly distributed over the entire dorsum (see Gosliner, Schaefer, & Miller, 1999), which never form aggregations as in Doriopsilla elitae. Doriopsilla tishae Valdés & Hamann, n. sp. (Figures 1D, 4—S, 3B) Doriopsilla sp. 3 — Valdés et al., 2006: 204—205. Material examined: HOLOTYPE: Coxen’s Hole, Roa- tan, Honduras, 23 December 1991, 1 specimen 49 mm long live (LACM 1932). PARATYPES: Coxen’s Hole, A. Valdés & J. Hamann, 2007 Page 213 Figure 3. Scanning electron micrographs of penial spines. A, Doriopsilla elitae (LACM 1931). B, Doriopsilla tishae (LACM 1933). Roatan, Honduras, 23 December 1991, 5 specimens 49 mm long live (LACM 1933); Soldado Channel, Guanaja, Honduras, 6 August 1991, 30 m depth, 2 specimens 25mm long live (LACM _ 1975); Little French Cay, Guanaja, Honduras, 10 August 1991, 1 specimen 27 mm long live (LACM 1976). External morphology: Living animals reach up to 49 mm in length. The general color of the living animals is translucent yellowish-white (Figure 1D). The dorsum is covered with a network of anastomosed, irregular, thick black lines running in between the dorsal tubercles. In addition, there are numerous minute opaque white spots concentrated near the edge of the mantle and on top of the dorsal tubercles, giving them the appearance of being completely white. The rhinophores and gill are pale yellow. The body is oval, low, stiffened by a subepidermal network of strong spicules over the entire body surface. The dorsum is covered by a number of large, Page 214 Table 1 The Veliger, Vol. 50, No. 3 Comparative table of the external differences among species of Doriopsilla in the Atlantic Ocean. Doriopsilla areolata Doriopsilla albolineata Doriopsilla nigrolineata Doriopsilla pelseneeri Doriopsilla pharpa Doriopsilla espinosal Doriopsilla elitae sp. n. Doriopsilla tishae sp. n. Body color Dorsal tubercles Yellow to pinkish, reddish or pale brown with white rings or lines forming a network, center of the dorsum darker Pearl gray with white lines most of which are transverse, brown lines in the mantle margin Translucent yellowish-gray with white rings around tubercles and black lines forming a network White, yellow, orange or red, with a white line around the gill pocket edge Yellow to orange with numerous dark brown spots on the whole surface of the dorsum Translucent white to yellowish with several large opaque white patches, and a number of conspicuous red spots Yellow to dark orange with opaque white patches situated on top of many dorsal tubercles Translucent yellowish-white with a network of thick black lines and numerous minute opaque white spots near the mantle edge and on the tubercles Low and rounded to conical tubercles, larger in two rows between rhinophores and gills Low and simply rounded tubercles, larger in two rows between rhinophores and gills Low and simply rounded tubercles, medial tubercles are higher and larger Large irregular warts, larger in the center of the dorsum Numerous and minute tubercles, all of them of a similar size Numerous low, simple conical tubercles, medial tubercles larger Low, simple, conical tubercles Large, semispherical tubercles, larger in the center of the dorsum Geographical range Eastern Atlantic: from northern Spain to the Cape Verde Islands and Mediterranean Sea. Western Atlantic: Virgin Islands, Puerto Rico, Martinique Atlantic coast of Africa, from Ghana to Angola Panama, Honduras Iberian Peninsula Atlantic coast of the continental USA from Maryland to Florida, and Cuba, Virgin Islands Cuba, Bahamas Aruba, St. Vincent and the Grenadines, Grenada, and Martinique Honduras semispherical tubercles, stiffened with spicules. Tuber- cles medial on the dorsum are larger, decreasing in size toward the borders of the mantle. The mantle margin is wide and_ slightly undulate. The rhinophores are perfoliate with up to 15 lamellae. The gill is composed of four tripinnate leaves. The anus 1s eccentric to the left. The oral tentacles are fused and grooved laterally. The anterior border of the foot is notched. Anatomy: The buccal bulb is elongate (Figures 4A, 5A), covered by minute, rather undifferentiated oral glands on its proximal portion. The tubular esophagus leads from the buccal bulb. At this point two retractor muscles insert onto the posterior of the bulb. The esophagus is very long and convoluted (Figure 5B). Posteriorly, it broadens into a large muscular portion. The intestine runs posteriorly in the usual position and lacks any pyloric gland. The ampulla is simple, oval (Figures 4C, 5B). It divides into a short oviduct, which enters the female gland and the prostate. The prostate is broad, flattened. From its distal end, the prostate leads into an elongated deferent duct. The penis, when everted, is very long and contains several rows of penial hooks. The penial hooks are approximately 15 um wide at the base and up to 20 um in length (Figure 3B). The vagina is very long and convoluted. At its proximal end 1s a large, thin-walled, spherical bursa copulatrix. The seminal receptacle is small, having a long duct that joins the vagina at the point where it connects the bursa copulatrix. From this point also emerges the uterine duct. The circulatory system consists in a large heart (Figures 4A, 5A), joined by the aorta with a flattened blood gland, situated behind the central nervous system. Etymology: The species is named after Tisha Thiessen, oldest daughter of Jeff Hamann. Geographic range: Thus far this species is only known from the Bay Islands, Honduras. Remarks: Doriopsilla tishae 1s most similar to Dor- iopsilla nigrolineata (Figure 1F), originally described from the Caribbean of Panama, due to the presence of dorsal black lines in both species. Differences between these two species include the external coloration and the anatomy. Externally, both species have a pattern of black lines on the dorsum, however, in D. tishae the lines are more conspicuous, thicker, longer and consistently anastomosed, whereas in D. nigrolineata A. Valdés & J. Hamann, 2007 Figure 4. Drawings of the internal anatomy of the paratype of Doriopsilla tishae (LACM 1933). A, General view of the anatomy, scale bar = | mm. B, Detail of the anterior portion of the digestive system, scale bar = 1 mm. C, Reproductive system, scale bar = 1mm. D, Connection of the bursa copulatrix and seminal receptacle, scale bar = 1 mm. Abbreviations: am, ampulla; bb, buccal bulb; bce, bursa copulatrix; bg, blood gland; dd, deferent duct; dg, digestive gland; e, esophagus; fg, female gland; ht, heart; i, intestine; pr, prostate; rm, retractor muscle; rs, reproductive system; s, syrinx; sr, seminal receptacle; v, vagina. they are generally shorter, thinner and rarely contact each other (Figure 1F). Additionally, the tubercles of D. tishae are much larger than those of D. nigrolineata, and are completely covered with opaque white spots, whereas in D. nigrolineata the white spots surround smaller and more elongate tubercles. For a comparison of the external characteristics of D. tishae with other Atlantic species of Doriopsilla see Table 1. Internally, the structure of the reproductive and digestive system is different. In D. nigrolineata the proximal region of the intestine is more inflated and the proximal region of the esophagus is proportionally smaller than those of D. tishae (see Figure 6). The buccal bulb of D. tishae is elongate, whereas that of D. nigrolineata is shorter. The seminal receptacle of D. nigrolineata is proportionally larger and joined to the bursa copulatrix by a much shorter duct than in D. tishae. No other Atlantic species of Doriopsilla have a network of dorsal black lines. There are no eastern Pacific species of Doriopsilla with a color pattern similar to that of Doriopsilla tishae. Doriopsilla areolata Bergh, 1880 (Figures 1E, 7) Material examined: Anse Noire, North Point, Martini- que, 15 July 1987, 30 m depth, 1 specimen 36 mm long Page 215 Figure 5. Drawings of the internal anatomy of a paratype of Doriopsilla tishae (LACM 1975). A, General view of the anatomy, scale bar = 1 mm. B, Reproductive system, scale bar = | mm. Abbreviations: am, ampulla; bb, buccal bulb; bce, bursa copulatrix; bg, blood gland; dd, deferent duct; dg, digestive gland; e, esophagus; fg, female gland; ht, heart; 1, intestine; pr, prostate; rm, retractor muscle; sr, seminal receptacle; v, vagina. live (LACM 173780). Additional specimens were photographed but are not preserved. External morphology: Living Caribbean animals of this species reach 36 mm in length. The general color of the Caribbean animals varies from yellowish to pinkish or reddish (Figure 1E; Valdés et al., 2006). The dorsum is completely covered with numerous minute opaque bluish-white dots. The largest tubercles, situated on the outer side of the dorsum and inner mantle margin as well as on the branchial sheath, are surrounded by conspicuous white rings composed of accumulations of opaque white dots. The rhinophores and gill are the same color as the dorsum. The body is oval, low, stiffened by a subepidermal network of strong spicules over the entire body surface. The dorsum is covered by a number of round to conical tubercles, stiffened with spicules. Tubercles on the sides of the dorsum are larger, decreasing in size toward the borders of the mantle and the center of the dorsum. The mantle margin is wide and slightly undulate. The Page 216 Figure 6. Drawings of the internal anatomy of Doriopsilla nigrolineata (LACM 173781). A, General view of the anatomy, scale bar = 1 mm. B, Reproductive system, scale bar as in C. C, Connection of the bursa copulatrix and seminal receptacle, scale bar = 1 mm. Abbreviations: am, ampulla; bb, buccal bulb; be, bursa copulatrix; bg, blood gland; dd, deferent duct; dg, digestive gland; e, esophagus; fg, female gland; ht, heart; 1, intestine; pr, prostate; rm, retractor muscle; rs, reproductive system; s, syrinx; sr, seminal receptacle; v, vagina. rhinophores are perfoliate with up to 12 lamellae. The gill is composed of five tripinnate leaves. The anus 1s eccentric to the left. The oral tentacles are fused and grooved laterally. The anterior border of the foot is notched. Anatomy: The buccal bulb is very elongate (Figure 7A— B), covered by minute, rather undifferentiated oral glands on its proximal portion. The tubular esophagus leads from the buccal bulb. The esophagus is very long and convoluted (Figure 7B). Posteriorly, it broadens into a small muscular portion. The intestine runs posteriorly in the usual position and lacks any pyloric gland. The ampulla is simple, very elongate (Figure 7D). It divides into a short oviduct, which enters the female gland and the prostate. The prostate is broad, flattened (Figure 7C). From its distal end, the prostate leads into a short deferent duct. The penis, when everted, is very long and contains several rows of penial hooks. The vagina is very long and convoluted particularly at its proximal end where it connects to the thin-walled, spherical bursa copulatrix. The seminal receptacle is small, having a long duct that joins the vagina at the point where it connects to the bursa copulatrix. The uterine duct also emerges from this point. ihe Veliger,- Vol) 505 Now < Figure 7. Drawings of the internal anatomy of Doriopsilla areolata (LACM 173780). A, General view of the anatomy, scale bar = | mm. B, Detail of the anterior portion of the digestive system, scale bar = 1 mm. C, Reproductive system, scale bar = | mm. D, Connection of the bursa copulatrix, seminal receptacle, and ampulla, scale bar = 1 mm. Abbre- viations: am, ampulla; bb, buccal bulb; be, bursa copulatrix; bg, blood gland; dd, deferent duct; dg, digestive gland; e, esophagus; fg, female gland; ht, heart; i, intestine; pr, prostate; rm, retractor muscle; rs, reproductive system; s, syrinx; sr, seminal receptacle; v, vagina. Remarks: Eastern Atlantic specimens of this species have been described in detail by Valdés & Ortea (1997). Caribbean specimens are anatomically identical to those from the eastern Atlantic, but they lack the pattern of irregular white lines on the dorsum. Instead they only display the white rings present in juvenile specimens from the eastern Atlantic. Marcus & Marcus (1962) cited Doriopsilla areolata for the Caribbean (St. John, Virgin Islands) based on a single specimen that was preserved before study, therefore the color of living Caribbean animals remained unknown. The anatomical descriptions by Marcus & Marcus (1962) match our observations and the descriptions for eastern Atlantic animals, confirm- ing that they are all members of the same species. Doriopsilla nigrolineata Meyer, 1977 (Figures 1F, 6) Material examined: Michael Rock Channel, Guanaja, Honduras, 5 August 1991, 2-3 m depth, 1 specimen 25 mm long, live (LACM 173781). Punta Hospital, Bocas del Toro, Panama, 23 August 2006, 10 m depth, 1 specimen 25 mm long live (LACM 173782). External morphology: Living animals reach up to A. Valdés & J. Hamann, 2007 25mm in length. The general color of the living animals is translucent yellowish-gray (Figure 1F). The viscera is visible through the skin as a dark gray area. The dorsum is covered with a series of thin, irregular black lines running in between the dorsal tubercles and almost never contacting each other. In some specimens the lines may be longer than in others. In addition, there are numerous minute opaque white spots all over the dorsum, forming rings around the dorsal tubercles but never on the tubercles themselves. The rhinophores and gill are pale yellow. The rhinophores have white apices. The body is oval, low, stiffened by a subepidermal network of strong spicules over the entire body surface. The dorsum is covered by a number of round tubercles, stiffened with spicules. Medial tubercles on the dorsum are higher and larger, decreasing in length and size toward the borders of the mantle. The mantle margin is wide and slightly undulate. The rhinophores are perfoliate with up to 12 lamellae. The gill is composed of four tripinnate leaves. The anus is eccentric to the left. The oral tentacles are fused and grooved laterally. The anterior border of the foot is notched. Anatomy: The buccal bulb is short and wide (Fig- ure 6A), covered by minute, rather undifferentiated oral glands. The tubular esophagus leads from the buccal bulb. The esophagus is short and convoluted. Posteriorly, it broadens into a muscular portion. The intestine 1s greatly expanded proximally, runs posteri- orly in the usual position and lacks any pyloric gland. The ampulla is simple, elongate (Figure 6B) and enters the female gland near the opening of the prostate. The prostate is broad and flattened. From its distal end, the prostate leads into a relatively short deferent duct. The penis, when everted, is very long and contains several rows of penial hooks. The vagina is short and convoluted. At its proximal end is an oval, thin-walled bursa copulatrix. The seminal receptacle is small, having a very short duct that joins the vagina at the point where it connects the bursa copulatrix (Figure 6C). From this point also emerges the uterine duct. The circulatory system consists in a large heart (Figure 6A), joined by the aorta with a flattened blood gland, situated behind the central nervous system. Geographic range: Originally described from Panama, this species has been subsequently collected and photographed from Honduras. Remarks: The original description of this species (Meyer, 1977) was based on external characteristics but did not include anatomical examinations. The redescription by Valdés & Ortea (1997) was a re- examination of the external morphology of the Page 217 holotype and produced no new information. The present paper provides the first anatomical examina- tion of this species, which clearly differs from Doriopsilla areolata in several regards. Differences include the length of the buccal bulb, which is several times longer in D. areolata than in Doriopsilla nigrolineata. Also the structure of the reproductive system is different in these two species, with a larger seminal receptacle in D. nigrolineata connected to the bursa copulatrix by a much shorter duct. Finally, the ampulla is proportionally longer in D. areolata. Externally these two species are readily distinguishable by the presence of irregular black lines in D. nigrolineata that are absent in D. areolata. DISCUSSION Cariopsillidae Ortea & Espinosa, 2005, type genus Cariopsilla Ortea & Espinosa, 2005, was described based on the species Doriopsilla pharpa Marcus, 1961, which has dorsal tubercles superficially similar to the caryophyllidia present in other dorid nudibranchs. Ortea & Espinosa (2005) indicated that this new taxon is proposed to rationalize the systematics of radula-less dorids, independently from phylogenetic studies that produce erroneous results due to the incorrect use of characters. More specifically, Ortea & Espinosa (2005) criticized the phylogenetic analysis conducted by Valdés & Gosliner (1999) because of the exclusion of characters referring to the caryophyllidia-like tubercles of D. pharpa. According to the only available phylogenetic hy- pothesis including D. pharpa (Valdés & Gosliner, 1999), this species is nested in the Doriopsilla clade, so the introduction of a new genus and a new family for this species would render both Doriopsilla and Dendrodor- ididae as paraphyletic. Ortea & Espinosa (2005) did not provide arguments to support the evolutionary distinc- tiveness of D. pharpa or a phylogenetic hypothesis to support their alternative grouping. For all these reasons, and until new evidence becomes available, we regard Cariopsillidae as a junior synonym of Dendrodorididae and Cariopsilla as a junior synonym of Doriopsilla. Acknowledgments. This paper has been supported by the US National Science Foundation through the PEET grant “Phylogenetic systematics of the Nudibranchia’” (DEB- 0329054) to T.M. Gosliner and the senior author. The SEM work was conducted at the Natural History Museum of Los Angeles County facility supported by the NSF MRI grant DBI-0216506. Lindsey Groves (LACM) curated the specimens collected and critically reviewed the manuscript. An anonymous reviewer and Elita Hamann made constructive comments on the manuscript. Elita Hamann also reviewed the manuscript for grammar and consistency. Page 218 REFERENCES GOSLINER, T. M., M. SCHAEFER & S. V. MILLER. 1999. A new species of Doriopsilla (Nudibranchia: Dendrodoridi- dae) from the Pacific Coast of North America, including a comparison with Doriopsilla albopunctata (Cooper, 1863). The Veliger 42:201-210. Marcus, Ev. & ER. MARCUS. 1962. Opisthobranchs from Florida and the Virgin Islands. Bulletin of Marine Science of the Gulf and Caribbean 12:450-488. OrTEA, J. & J. ESPINOSA. 2005. Nueva familia y nuevo género de Doridos Porostomados. Avicennia 18:64—-66. VALDES, A. & T. M. GOSLINER. 1999. Phylogenetic system- atics of the radula-less dorids (Mollusca: Nudibranchia) with the description of a new genus and a new family. Zoologica Scripta 28:315—360. VALDES, A. & T. M. GOSLINER. 2001. Systematics and ithe Veliger, Vol, 50 sNoms phylogeny of the caryophyllidia-bearing dorids (Mollus- ca, Nudibranchia), with the description of a new genus and four new species from Indo-Pacific deep waters. Zoological Journal of the Linnean Society 133:103— 198. VALDES, A. & J. ORTEA. 1997. Review of the genus Doriopsilla Bergh, 1880 (Gastropoda: Nudibranchia) in the Atlantic Ocean. The Veliger 40:240-254. VALDES, A. & J. ORTEA. 1998. A new species of Doriopsilla (Mollusca, Nudibranchia, Dendrodorididae) from Cuba. Proceedings of the California Academy of Sciences 50: 389-396. VALDES, A., J. HAMANN, D. W. BEHRENS & A. DUPONT. 2006. Caribbean Sea Slugs: A Field Guide to the Opisthobranch Mollusks from the Tropical Northwestern Atlantic. Sea Challengers: Gig Harbor, Washington. 289 pp. The Veliger 50(3):219—224 (October 1, 2008) THE VELIGER © CMS, Inc., 2007 Dendropoma mejillonensis sp. nov., a New Species of Vermetid (Caenogastropoda) from Northern Chile ALDO PACHECO!* AND JURGEN LAUDIEN? ‘Universidad de Antofagasta, Facultad de Recursos del Mar, Av. Angamos 601, PO Box. 117, Antofagasta, Chile ° Alfred Wegener Institute for Polar and Marine Research, D-27568 Bremerhaven, Germany Abstract. _Dendropoma mejillonensis sp. nov. is described based on morphology for the first time. This vermetid gastropod inhabits the rocky subtidal zone of Peninsula Mejillones in northern Chile. In July 2006, specimens were collected by SCUBA divers from the rocky ““Anemones Wall” (23°28'17.30"S, 70°37'13.80”W) at 17 m depth. The morphology of D. mejillonensis is distinguished from that of other members of the family by its pointed lip on the external border of the protoconch and the two white bands on the head tentacles. This extends the geographical range of the genus Dendropoma into the Southeastern Pacific. The present species D. mejillonensis is the only known vermetid gastropod able to thrive under the cold upwelling conditions of the Humboldt Current ecosystem off northern Chile. INTRODUCTION Marine gastropods of the family Vermetidae are sessile organisms with an irregular, uncoiled shell providing a three-dimensional biogenic habitat for associated species. Their distribution is restricted to tropical and subtropical latitudes (Mexico, California and West Africa) as well as to locations in the warm Mediterra- nean Sea (Keen, 1961, 1971; Schiaparelli et al., 2003). Habitats are rocky intertidal and subtidal zones with warm and oxygenated waters (Keen, 1961; Calvo et al., 1998). Due to the irregular tube form, taxonomic identification has commonly been confused with Vermicularia (Turritellidae) (Bieler, 1996) and Serpulid polychaetes (Keen, 1961, 1971) resulting in a confused taxonomic status. The morphological characters deemed useful for taxonomic identification have changed over time (Bieler, 1995; Schiaparelli & Métivier, 2000). A genetic study further concluded that disjunct populations of Dendropoma species are close phylogenetic relatives (Rawlings et al., 2001), thus suggesting that taxonomic determination should be approached carefully. The genus Dendropoma (Morch, 1861) was reviewed by Keen (1961) on the basis of 10 species distributed among tropical and subtropical locations. Distinctive morphological characteristics for this genus are pla- norboid early whorls that become more loosely coiled in later stages; and the sculpture of lamellar growth- striations that may or may not be intersected by longitudinal lines, sinuous and rising toward a crest near the outer edge of the whorl in most species. The * Tel.: 056-55-637404, Email: babuchapv@yahoo.com operculum is well developed and equal in diameter to the aperture. At present, the genus Dendropoma covers intertidal and sublittoral species and can be gregarious or solitary. So far, the most comprehensive information about Dendropoma spp. taxonomy is provided by Hadfield et al. (1972) for specimens found off Hawaii. Information on the distribution of vermetids off continental Chile and its offshore islands is scarce and the taxonomic status is still uncertain (Rehder, 1980: Ramirez & Osorio, 2000; R. Bieler pers. comm.). In fact, extensive reviews of gastropod taxonomy and studies of invertebrate biogeographic patterns available from this coast do not mention the family in the region (Marincovich, 1973; Guzman et al., 1998; Brattstrom & Johanssen, 1983; Valdovinos, 1999; Lancellotti & Vasquez, 2000). Anecdotally, vermetids have been observed associated with holdfasts of the kelp Lessonia trabeculata Villouta & Santelices, 1986 off central Chile (Vasquez & Vega, 2004). With the exception of the latter observation, there is no published evidence from the Chilean coast. Nonetheless, Dendropoma platypus Morch, 1861; Dendropoma spp. and Serpulorbis Sassi, 1827 have been recorded from Easter Island (Rehder, 1980; Ramirez, 1987; Valdovinos, 1999) and Serpulor- bis sp. was also observed at Robinson Crusoe Island (Juan Fernandez archipelago) (Ramirez & Osorio, 2000), both insular Chilean locations. Northern Chile forms part of the Humboldt Current upwelling ecosystem, which is characterized by year- round high levels of primary production due to wind- driven cold upwelling water, which returns nutrients to the euphotic zone (Barber & Smith, 1981). There is a shallow oxygen minimum zone (OMZ) and only the upper 40 m are well oxygenated (Arntz et al., 2006). Page 220 Peninsula Mejillones Isla Santa Maria Antofagasta bay Pacific Ocean A 170° 37' 13,80" the Veliger Vols 50) Noms Figure 1. A. Sampling location “‘Anemones Wall” (23°28'17.30"S, 70°37'13.80”W) opposite the southeastern side of Isla Santa Maria. B. Peninsula Mejillones. C. Distributional range of related vermetid species (a) Petalonchus innumerabilis (b) Serpulorbis squamigerus (c) Vermetus compta (d) Dendropoma rastrum (e) Serpulorbis sp. (f) Dendropoma platypus. This habitat is very different from that of warm-water subtropical and tropical vermetid species. In this study Dendropoma mejillonensis sp. nov. is described from Peninsula Mejillones, a location within this particular upwelling system. A detailed morphological character- ization is provided. MATERIAL AND METHODS Individuals of Dendropoma mejillonensis sp. nov. colonizing a vertical rock wall in the subtidal zone (17 m depth) of Peninsula Mejillones (23°28'17.30"S, 70°37'13.80"W) were photographed and collected by SCUBA divers on July 11th, 2006 (Figure 1A, B). Several vermetid clusters were scratched from the rock with a knife and maintained in the laboratory for observations. Measurements were taken with a digital caliper or by using calibrated eyepieces on a dissecting microscope. Photographs were taken with a Canon Power Shot S50 camera connected to a_ binocular microscope Olympus SZ61. Animals were anesthetized by adding methanol drops in the small examination containers before sacrificing. Soft bodies were removed from the shell after cracking with a small clamp. Gross anatomy of the soft parts was studied under a dissection microscope. Air-dried shells, radula, proto- conch and opercula were observed and photographed, using the scanning electron microscope JEOL, model JSM- 6360LV. Diagnosis Genus Dendropoma Morch 1861 Solitary to colonial forms, corroding a trench in the substrate, in which the lower part of each volution is embedded; coiling planorboid in early whorls, becom- ing looser in later whorls, with tendency toward right- angle turns. The color of the adult is mostly white, intermittently stained with dark brown, especially within. The sculpture of lamellar growth-striations, that may or may not be intersected by longitudinal lines, is sinuous and rises toward a crest near the outer edge of the whorl in most species. Two nuclear whorls are dark brown in color, inflated, smooth to malleated or axially ribbed, and the aperture lip is pointed or claw-like in some species. The operculum is well developed, as large as the aperture, its inner surface having a distinct central attachment scar that is somewhat button-like, and its exterior composed of chitinous plates in a spiral arrangement, either com- pactly welded to form a smooth surface or variously agglutinated with foreign materials. Dendropoma mejillonensis sp. nov. Type locality: Live-taken syntypes collected from a large aggregation colonizing Anemones wall at 17 m depth, Peninsula Mejillones, northern Chile (23°28'17.30"S, 70°37'13.80"W) were deposited in the Field Museum of Natural History, Chicago, Illinois, U.S.A. (FMNH N°- 312172 and N°-312173). Additional samples were deposited in the Museo Nacional de Historia Natural de Santiago de Chile (paratype MNHNCL N°-5159 and syntypes MNHNCL N°-5160, 5161, 5162) Teleoconch (Figure 2a, b): The tubes form continuous and compact colonies, which are grey to faintly green in the field, but white after cleaning. Jn situ, the tubes are slightly nested in the rocky substrate. The attached part of the tube appears eroded, and thus is thinner (Figure 2d). The aperture is circular and its mean A. Pacheco & J. Laudien, 2007 Page 221 Figure 2. Dendropoma mejillonensis sp. nov. (a) Mass of living adult tubes. (b) Lateral view (c) Teleconch of juvenile showing concentric growth striations. (d) Smooth eroded part showed from the attached part of the tubes. (e) Operculum dorsal view. (f) Operculum ventral view. (g) Protoconch aperture. (h) Protoconch; detail of the sculpture and pointed lip-like external border. (1) Protoconch in ventral position, the earliest whorl is noted. (j) Radula displacement. (k) Detail of the cusp on the marginal teeth. (1) View of the animal head, the arrow points to the distinctive white mark. diameter in adults is 4.29 mm (SD = 0.37; n = 16). The tube exhibits sinuous growth lines and the sculpture of lamellar growth-striations is not intersected by longi- tudinal lines (Figure 2c). The periostracum is white and the intermediate layer slightly cream. Observing from a cross-longitudinal section, three layers of the conch are present. The interior part is cream porcelain, darker towards the interior tube. Very soft longitudinal lines are only observed under magnification. There is no internal shell lamellar structure. The proximal part of the tube slightly tends to vertically rise from the rest of the mat. The coiling pattern is variable. Early whorls are like Planorbidae, coiling counterclockwise, fol- lowed by a very loose coiling or irregular pattern. The shell of the juvenile is white and translucent with clear axial ribs (Figure 2c). Operculum (Figure 2e, f): The form is circular and concave, slightly flattened and reddish in the center, brown-orange to colorless towards the external border. The diameter is 2.7 mm (SD = 0.2; n = 10) in adult specimens and about 1/5 of the length of the relaxed Page 222 pedal disk diameter. The operculum is composed of concentric layers of chitinous material with visible concentric irregular lines, notably in juveniles. The small mamilla is inserted in the pedal surface. Almost 90% of the studied opercula were fouled with bryozoans. Protoconch (Figure 2g, h, i): Globular, brown or colorless, white towards the earliest whorl. The shell shows 1 to 1.5 nuclear whorls, ornamented with longitudinal grooves. The grooves show no evident axial pattern, are variable in size and present a slightly rectangular or triangular shape with no marks at the corners. The external border presents a pointed lip shape and growths striations are present. At hatching, shell length (the distance from the external lip border to the opposite whorl margin) is 0.77 mm (SD = 0.07; n = 10). Radula (Figure 2j, k): Taenioglossan type, similar to the description of other vermetids (i.e., Vermetus triquetrus Bivona-Bernardi, 1832 and Thylaeodus rugu- losus Monterosato, 1878; Bieler, 1995), transparent, consisting on average of 39.8 (SD = 6.06) rows of teeth (counts and measurements based on adult animals of 4 mm shell aperture, n = 10, no differences between sexes were noted). Total length of radular ribbon 1s 2.35 mm (SD = 0.34) and 0.196 mm width (SD = 0.011, mid ribbon). A trapezoidal rachidian tooth with a strong main cusp and 4—5 flanking cups on either side (diminishing toward margin), basal denticles strongly developed. Lateral tooth cusp arrangement of triangu- lar cutting shape, as in the central tooth, with two flanking cusps on either side. The inner marginal tooth is slender with a strong main cusp and the inner marginal with one flanking cusp on inside and two on outside. The outer marginal teeth present a single flanking cusp smooth on outside. Radular formula: 2+1+R+1+2. Animal: Removed from the shell the body is short and narrowest towards the terminal part, which is slightly coiled. The average length of relaxed large adult specimens is 18.66 mm (SD = 1.68; n = 10). The head is mainly light grey or reddish with black, white and yellow specks. The posterior part is reddish or dark brown 1n color. Two white bands on the head tentacles are distinctive appearing as a white eyebrow (Fig- ure 21). The head tentacles are brown or light grey in color with black and yellow dots, no distinctive marks at the tips are visible. The pedal tentacles are light grey with yellow specks. In both sexes the light orange/ melon mantle is entire and is characterized by a light brown border. The foot is a similar color to the mantle; however it has a white band around the operculum insertion. The gill filaments are about 1/3 of the size of the mantle and slightly triangular in shape. The The Veliger, Vol. 50, No. 3 Figure 3. Capsules of Dendropoma mejillonensis sp. nov. containing juveniles (arrow left side) and nurse yolk (arrows right side). columellar muscle appears as a white triangular narrow strip, enabling the animal to retreat deeply into its shell. Female’s broods comprise three to four egg capsules, which are ovoid and the membrane is translucent. Each capsule contains between three to ten juveniles. Early capsules contain nurse yolk (Figure 3). Feeding is carried out by mucous threads. Habitat: The specimens were attached to a vertical rock wall, which extends from the shallow subtidal down to 50 m depth. In the field, colonies showed a light grey to white color and were commonly fouled by calcareous algae causing a red/purple coloration. The surrounding benthic community is dominated by the kelp Lessonia trabeculata from 13 m depth down to 25 m. Below 25m, kelp abundance is substantially reduced and relatively small epibenthic taxa such as calcareous algae (Lythothamniom sp. and Lithophyllum sp.), red algae (Rhodymenia corallina Bory de Saint Vincent & Greville), bryozoans (Membranipora isabelleana D’Or- bigny, 1847 and Lagenicella variabilis Moyano, 1991), and Porifera cover the substrate. Dendropoma mejillo- nensis sp. nov. colonies were observed between 15 and 25 m. Etymology: The species is named Dendropoma mejillo- nensis in reference to the discovery location Peninsula Mejillones. DISCUSSION Taxonomic remarks The morphological classification of the species to the Dendropoma genus was carried out following Keen (1961). Dendropoma_ mejillonensis sp. nov. shows similarities to Dendropoma gregaria Hadfield & Kay, A. Pacheco & J. Laudien, 2007 1972 (Hadfield et al., 1972) from Hawaii, sharing the circular pattern in the operculum and dense white pigmentation around the eyes. The most noteworthy difference is in the protoconch sculpture, while D. gregaria has light axial ribs crossed by finer spiral striations, D. mejillonensis shows soft grooves without evident design shape and pattern. Distributional Remarks As already mentioned, the presence of vermetid gastropods is limited along the Pacific coast of South America. Alamo & Valdivieso (1997) reported Petalo- conchus innumerabilis Pilsbry & Olsson 1935 from Mazatlan (Mexico) to Bocapan and Huacho (Peru), Serpulorbis squamigerus Carpenter, 1857 from San Diego (California) to Paita (Peru) and Vermetus compta Carpenter, 1857 from British Columbia (Ca- nada) to Paita. Keen (1971) recorded Dendropoma lituella Morch, 1861 and Dendropoma rastrum Morch, 1861 from the northern part of the Eastern Pacific; both were found from southern California to the southern Gulf of California at La Paz, Baja California (see also Figure Ic). The presence of Dendropoma mejillonensis in the rocky subtidal zone of Peninsula Mejillones clearly extends the geographic range of the family into the Southeastern Pacific, almost 2000 km southwards. According to the literature the closest distribution limit of vermetids is Huacho (11°6'56.21”S, 77°37'9.46"W) (Alamo & Valdivieso, 1997). Easter Island and Juan Fernandez may be source locations if Dendropoma sp. is D. mejillonensis, in this case the range would be extended from insular to continental Chile. However, it is not possible to define the biogeography of this species, as we did not sample south or north of the type locality. Our record provides evidence that D. mejillonensis is able to thrive under cold upwelling conditions. The observed recruitment at Anemones Wall (A. Pacheco unpublished data) indicates that this species has the capacity to adapt to cold upwelling conditions. The species’ distribution may be limited by the presence of a short larval stage. As in the case of many other vermetids (Keen, 1961; Hadfield et al., 1972; Calvo et al., 1998), larvae of D. mejillonensis leave the female mantle cavity well developed and crawl around for less than one hour before cementing themselves to the substrate. The recent discovery from Peninsula Mejil- lones suggests that several unexplored areas with unreported species may still exist along the northern Chilean coastline, particularly in zones difficult to reach (Camus, 2001). Furthermore, distributions of rafting species (a dispersal mechanism suggested for vermetids (Bieler, 1995)) may extend quickly with an increasing amount of anthropogenic floating material, Page 223 facilitating the supply of sessile species to new regions (Thiel & Haye, 2006). A genetic study is necessary to reveal linkages between D. mejillonensis and other vermetids. Acknowledgments. We are grateful to Christian Guerra and Ivan Marin for their support during diving. Martha Calvo provided useful literature. José Riascos, Olaf Heilmayer, Ricardo Guinez are thanked for their suggestions on an earlier version of the manuscript. Carrie Auld and Ruth Alheit kindly edited the English. Pabla Vega helped with the SEM photographs. Rudiger Bieler provided taxonomic advice. We appreciated the comments made by two anonymous reviewers. This study was conducted in the framework of the EU-funded INCO project, “Climate Variability and El Nino-Southern Oscillation: Implications for Natural Coastal Resources and Management” (CENSOR) and Programa Bicentenario de Ciencia y Tecnologia RUE-02. This is CENSOR publication N° 341. EMERATURE, CIbED ALAMO, V. & V. VALDIVIESO. 1997. Lista sistematica de moluscos marinos del Peru. Instituto del Mar del Pert, 183 pp. ARNTZ, W., V. GALLARDO, D. GUTIERREZ, E. ISLA, L. LEVIN, J. MENDO, C. NEIRA, G. T. ROWE, J. TARAZONA & M. WOLFF. 2006. El Nino and similar perturbation effects on the benthos of the Humboldt, California and Benguela current upwelling ecosystems. Advances in Geoscience 6:243—265. BARBER, R. T. & R. L. SMITH. 1981. Coastal Upwelling Ecosystems. In: A. R. Longhurst (ed.), Analysis of Marine Ecosystems. Academic Press. Pp. 31—68. BIELER, R. 1995. Vermetids gastropods from Sao Miguel, Azores: Comparative anatomy, systematic position and biogeographic affiliation. Acoreana, Supplement. 173— 192. BIELER, R. 1996. Morch’s worm snail taxa (Caenogastropoda: Vermetidae, Siliquariidae, Turritellidae). American Ma- lacological Bulletin 13(1/2):23-35. BRATTSTROM, H. & A. JOHANSSEN. 1983. Ecological and regional zoogeography of the marine benthic fauna of Chile. Sarsia 68:289—339. CALvo, M., J. TEMPLADO & P. E. PENCHASZADEH. 1998. Reproductive biology of the gregarious mediterranean gastropod Dendropoma petraeum. Journal of the Marine Biological Association of the United Kingdom 78:525— 549. Camus, P. A. 2001. Marine biogeography of continental Chile. Revista Chilena de Historia Natural 74:587-617. GUZMAN, N., S. SAA & L. ORTLIEB. 1998. Descriptive catalogue of nearshore molluscs (Gastropoda and Pele- cypoda) from Antofagasta area, 23°S (Chile). Estudios Oceanologicos 17:17-86. HADFIELD, M. G., E. A. KAy, M. U. GILLETTE & M. C. LLOYD. 1972. The Vermetidae (Mollusca: Gastropoda) of the Hawaiian Islands. Marine Biology 12:81—98. KEEN, A. M. 1961. A proposed reclassification of the gastropod family Vermetidae. Bulletin of the British Museum (Natural History) Zoology 7(3):181-213. KEEN, A. M. 1971. Sea shells of tropical west America. Marine mollusks from Baja California to Peru. Stanford University Press: Stanford, California. 1063 pp. Page 224 LANCELLOTTI, D. A. & J. A. VASQUEZ. 2000. Zoogeography of benthic macroinvertebrates of the Chilean coast: contribution for marine conservation. Revista Chilena de Historia Natural 73:99-129. MARINCOVICH, L. JR. 1973. Intertidal mollusks of Iquique, Chile. Natural History Museum Los Angeles County Science Bulletin 16:175—226. RAMIREZ, J. 1987. Moluscos de Chile. II. Mesogastropoda. Santiago de Chile Imprenta Museo Nacional de Historia Natural: Chile. 172 pp. RAMIREZ, M. E. & C. OsoRIO. 2000. Patrones de distribucion de macroalgas y macroinvertebrados intermareales de la isla Robinson Crusoe, archipiélago de Juan Fernandez, Chile. Investigaciones Marinas Valparaiso 28:1—13. RAWLINGS, T., T. COLLINS & R. BIELER. 2001. A major mitochondrial gene rearrangement among closely related species. Molecular Biology and Evolution 18:1604—1609. REHDER, H. A. 1980. The marine mollusks of Easter Island (Isla de Pascua) and Sala y Gomez. Smithsonian Contributions to Zoology N°- 289. SCHIAPARELLI, S. & B. METIVIER. 2000. On the identity of The Veliger, Vol. 50, No. 3 “Vermetus” roussai Vaillant 1871 (Mollusca, Caenogas- tropoda, Vermetidae), with description of the new species. Zoosystema 22:677—687. SCHIAPARELLI, S., P. GUIDETTI & R. CATTANEO-VIETTI. 2003. Can mineralogical features affect the distribution patterns of sessile gastropods? The Vermetidae case in the Mediterranean Sea. Journal of the Marine Biological Association of the United Kingdom 83:1267—1268. THIEL, M. & P. HAYE. 2006. The ecology of rafting in the marine environment III. Biogeographical and evolution- ary consequences. Oceanography and Marine Biology: An Annual Review 44:323-429. VALDOVINOS, C. 1999. Biodiversidad de moluscos chilenos: base de datos taxondmica y distribucional. Gayana 63: 111-164. VASQUEZ, J. A. & J. M. A. VEGA. 2004. Ecosistemas marinos costeros del Parque Nacional Bosque Fray Jorge. In: F. A. Squeo, J. R. Gutiérrez & L. R. Hernandez (eds.), Historia Natural del Parque Nacional Bosque Fray Jorge. Ediciones Universidad de La Serena: La Serena, Chile. 13:Pp. 235-252. THE VELIGER © 9) The Veliger 50(3):225-240 (October 1, 2008) © CMS, Inc., 2007 New Genera and Species of Peristerniinae (Gastropoda: Fasciolaridae) from the Caribbean Region, with Comments on the Fasciolariid Fauna of Bermuda WILLIAM G. LYONS 4227 Porpoise Drive SE, St. Petersburg, FL 33705, USA AND MARTIN AVERY SNYDER Department of Malacology, Academy of Natural Sciences of Philadelphia, 19'° and Benjamin Franklin Parkway, Philadelphia, PA 19103-1195, USA (e-mail: dr.martin.snyder@gmail.com) Abstract. Three new genera of Latirus-like gastropods from the western Atlantic are described and distinguished from Hemipolygona, Polygona, and Pustulatirus, all peristerniine genera with sympatric species in the region, and from Fusolatirus of the Indo-west Pacific. The new genera are: Lamellilatirus, type species Fusus ceramidus Dall, 1889a, formerly classified in Fusinus, Recent, southern Caribbean Sea; Lightbournus, type species L. russjenseni sp. nov., Recent, Bermuda; and Bullockus, type species B. guesti, Recent, Bermuda; Bullockus pseudovarai sp. nov., Recent, eastern Bahamas, is also described. Other species reclassified in Bullockus are Latirus (Hemipolygona) mecmurrayi Clench and Aguayo, 1941, from northern Cuba and the Bahamas; Latirus ( Latirus) varai Bullock, 1970, from eastern Cuba; and Hemipolygona honkeri Snyder, 2006, from the Bahamas and southwestern Caribbean Sea. Species of Bullockus generally live in upper slope depths (183-550 m), although one occurrence of B. guesti from 51 m is known. Pleistocene and Recent records of Latirus brevicaudatus (Reeve, 1847) at Bermuda are disproved; Pleistocene occurrence of Leucozonia nassa (Gmelin, 1791) at Bermuda is confirmed; and Pleistocene and Recent records of Fasciolaria spp. at Bermuda remain unconfirmed. INTRODUCTION not been addressed in those studies is Fusuws ceramidus Dall, 1889a. This species has usually been classified in Western Atlantic species of the fasciolariid subfamily the genus Fusinus Rafinesque, 1815 (e.g., Hadorn and Peristerniinae customarily have been classified in only Rogers, 2000), but Sunderland and Sunderland (1995) two genera, Latirus Montfort, 1810, and Leucozonia proposed that the shell seemed more like that of Gray, 1847 (e.g., Bullock, 1974; Vermeij and Snyder, Latirus, and Snyder (2003) noted that a review of 2002; Mallard and Robin, 2005). However, recent western Atlantic Latirus by Bullock (1968; unpublished studies of peristerniine taxa world-wide have reclassi- M.Sci. thesis) had indeed placed the species in that fied many species formerly in Latirus to other genera genus. Bullock’s account confirmed that the species is such as Benimakia Habe, 1958 (Vermeij and Snyder, peristerniine and, by criteria in use at that time, was 2003) and Fusolatirus Kuroda and Habe, 1971 (Snyder appropriately placed in Latirus, but it is excluded from and Callomon, 2005; Snyder and Bouchet, 2006). that genus by contemporary criteria. Vermeljy and Snyder (2006) further restricted Latirus To ascertain a proper generic assignment for to a relatively few Indo-west Pacific species, raised “Latirus’”’ ceramidus, we examined shells from the type Polygona Schumacher, 1817 and Hemipolygona Rover- locality, Barbados, and from another reported popu- eto, 1899, both formerly considered subgenera of lation at Bermuda. This examination convinced us that Latirus, to full generic rank, and introduced the new no generic name currently in use is appropriate for this generic names Pustulatirus and Turrilatirus to accom- species, so we introduce a new generic name for it. modate other species formerly classified in Latirus. “Latirus”’ ceramidus is redescribed, and evidence These studies prompted reassignment of nearly all pertinent to its generic distinctiveness is discussed. western Atlantic species formerly assigned to Latirus to The Bermuda material was found not to be “‘L.” Polygona, Hemipolygona, Benimakia, or Pustulatirus, ceramidus but rather to consist of two previously most by Vermeij and Snyder (2006). undescribed species which we also describe. These One western Atlantic species whose classification has species, both apparently endemic to Bermuda, are Page 226 designated as type species of two other new genera, one containing several additional Caribbean species, in- cluding another newly described here. The other new genus is monotypic. It is possible that these genera are deep-water derivatives of Hemipolygona. METHODS Specimens were examined from several institutional and private collections, identified by prefixes of catalogue numbers or collectors’ initials (see Abbrevi- ations). Shells were measured to the nearest 0.1 mm using vernier calipers. Unless otherwise stated, reported sizes are shell height (greatest length). Abbreviations ANSP ~— Academy of Natural Sciences of Philadelphia, PA. BMSM - Bailey-Matthews Shell Museum, Sanibel, FL. DMNH — Delaware Museum of Natural History, Wilmington, DE. FLMNH — Florida Museum of Natural History, Gainesville, FL. HGL — Collection of Harry G. Lee, Jacksonville, FL. KLS — Collection of Kevan & Linda Sunderland, Sunrise, FL. MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, MA. RH — Collection of Roland Hadorn, Lyss, Switzerland. Sh — empty (dead) shell. USNM — National Museum of Natural History, Smithsonian Institution, Washington, D.C. WGL — Collection of William G. Lyons, St. Peters- burg, FL. SYSTEMATICS Class Gastropoda Cuvier, 1795 Family Fasciolariidae J. E. Gray, 1853 Subfamily Peristerniinae Tryon, 1880 Lamellilatirus gen. nov. Type species: Fusus ceramidus Dall, 1889a, Recent, Barbados, designated herein. Diagnosis: Peristerniine gastropods with fusiform shells of small to medium size (adult lengths to 51.0 mm); whorls sculpted with moderate to strong axial ribs and less prominent spiral cords; sutures distinct, bordered anteriorly by prominent, dense band of lamellae; siphonal canals relatively short, slender, canted to left; aperture ovate, constricted anteriorly and posteriorly, with parietal shield bearing very weak, oblique columellar plicae and outer lip bearing internal lirae that are entire posteriorly but interrupted as beaded The Veliger, Vol, 50) Noms dots and dashes (sensu Vermeij and Snyder, 2006) anteriorly on mature specimens; radula of Latirus-type (see Cernohorsky, 1972:156-159 for examples of Latirus-type and Peristernia-type radulae). Etymology: Lamellilatirus, masculine, is a compound word formed of J/amella, Latin, the diminutive of lamina, or plate, in reference to the prominent subsutural lamellae of the shells, and the stem name Latirus, to acknowledge the place of the genus among the “‘Latirus-like”’ taxa. Remarks: Lamellilatirus is distinguished from Fusinus, where the type species was previously placed, by its peristerniine rather than fusinine radula (see Figure 3) and by having faint but definite oblique columellar folds (noted by Bullock (1968: 60)); shells of Fusinus lack columellar folds. Its relatively light-weight shells bearing conspicuous subsutural axial lamellae and faint oblique columellar plicae distinguish Lamellilatirus from other Latirus-like genera, most of which have heavier shells with well-developed, near-perpendicular columellar plicae. Shells of some species of the Indo- west Pacific genus Fusolatirus resemble those of Lamellilatirus but differ by having a Peristernia-like radula (see Snyder and Bouchet (2006: fig. 3k) for the radula of Fusolatirus). Abbott (1974) incorrectly classified Fusinus cerami- dus, the type species of Lamellilatirus, in the subgenus Barbarofusus Grabau and Shimer, 1909, which has been considered a subgenus or synonym of Fusinus or of Heilprinia Grabau, 1904. Shells of Barbarofusus lack columellar folds and subsutural lamellae, and their protoconchs are prominently ribbed on all whorls, whereas the protoconch of F. ceramidus is essentially smooth except for a few fine riblets near the junction with the teleoconch. Bullock (1968) and Sunderland and Sunderland (1995) proposed that Fusus ceramidus is more appropriately classified in Latirus, and Bullock (1968) proposed a manuscript name for a subgenus of Latirus with ceramidus as its type species. We chose not to validate that name because Bullock intended it collectively to represent several species that we do not believe represent a natural species-grouping. Lamellilatirus ceramidus (Dall, 1889a) Figures 1—2 Fusus ceramidus Dall, 1889a: 14, 171; Dall, 1890: 318, 359, pl. 6, fig. 6; Grabau, 1904: 74, 75; Lewis, 1965: 1067; Boss et al., 1968: 70; Bullock, 1968: 59, 106, 107, pl. 8, fig.7; Hadorn & Rogers, 2000: 14; Snyder, 2003: 64. Latirus ceramidus: Bullock, 1968: 59-61, 96, 106, pl. 3, figs. 5, 8. Fusinus ceramidus: Abbott, 1974: 230, 231, text-fig. 2530; Lyons, 1978: 87; Abbott & Dance, 1982: 189, W. G. Lyons & M. A. Snyder, 2007 row I, right fig. (4); Sander & Lalli, 1982: 313, 316, fig. 2; Snyder, 1984: 28, 30; Habe & Okutani, 1985: 193, row 1, right fig. (4); Sunderland & Sunderland, 1995: 18, 2 figs.; Goto & Poppe, 1996: 388; Hadorn, 1996: 18, 23, 24, fig. 1; Hadorn, 1997: 14; Hadorn & Rogers, 2000: 14, 39, 52, pl. 4, figs. 40, 41; Mallard & Robin, 2005: 11, pl. 18; [non Fusinus ceramidus (Dall, 1889), auctt., Bermuda, = Lightbournus russjenseni n. sp.]. Fusinus ( Barbarofusus) ceramidus: Abbott, 1974: 230. Fusinus caramidus [sic] “(Dall)”: Santos Galindo, 1977: 188; Snyder, 2003: 61. Fusinus ceraunicus [sic] “(Dall)”: Santos Galindo, 1977: 188; Snyder, 2003: 64. Fusinus ceramidas [sic]: Okutani, 1983: 24. Types examined: Lectotype, 46.2 mm, with operculum, Blake stn 290, Barbados, 13°11'54”N, 59°38'45’W, depth 134 m, USNM 87069; 2 paralectotypes, 18.7 & 11.3mm, Blake stn 273, Barbados, 13°03’05’N, 59°36'18”W, 188 m, USNM 87068. Other material examined: 5 sh, 39.0, 31.2, 30.0, 30.0 & 24.3 mm, in front of Bellair Research Institute, St. James coast, Barbados, depth 220 m, ANSP 416323; 6 sh, 51.0, 36.1, 31.3, 26.3, 19.4 & 19.2 mm, in front of Bellair Research Institute, St. James coast, Barbados, ANSP 416324; 4 sh, 43.7, 40.1, 39.7 & 35.3 mm, off west coast of Barbados, depth 165 m, dredged, ANSP 416371; 2 sh, 36.4 & 14.5 mm, west coast of Barbados, depth 90 m, WGL; 1 sh, 17.6 mm, west of Barbados, depth 166m, WGL; 1 sh, 31.1 mm, off Barbados, depth 202 m, dredged, WGL. Type locality: Barbados, 13°11'54"N, 59°38'45"W, depth 134 m. Description: Shell broadly fusiform, color pale orange- pink to white, length to 51.0 mm, with about 9-10 whorls. Protoconch of about 2 whorls, tip incurved and flat, sides convex but not expanding, first 1—3/4 whorls smooth, glassy, final 1/4 whorl with 2-4 axial riblets, junction with teleoconch distinct. Teleoconch with as many as 8 rounded, subtabulate, rapidly expanding whorls ornamented with axial ribs, spiral cords, and subsutural lamellae. Suture well defined by convexity of surrounding whorls, undulating slightly in accord with adjacent axial ribs and intercostal areas, bordered anteriorly by prominent, densely imbricated axial lamellae beginning on about third teleoconch whorl and continuing to anterior end of body whorl. Axial ribs prominent, broad, extending from suture to suture on first 2-3 whorls, beginning anterior to subsutural lamellae on subsequent whorls; usually 6 ribs on all whorls, less commonly 7 or 8 on penultimate and body whorls of some shells. Spiral cords generally low, broadest atop axial ribs, narrowest near centers of Page 227 intercostal areas; first three whorls with 2—3 primary cords crossing axial ribs and 3-4 fine threads between cords; primary cords increasing by intercalation to 5 by about whorl 6, third cord strongest, creating shoulder angle on that and subsequent whorls, posterior-most cord weaker than others; about 6 primary cords on penultimate whorl, secondary threads by now weak- ened and barely perceptible; body whorl subquadrate, defined by shapes of large axial ribs, ribs not continuing onto siphonal process; about 9-11 spiral cords on body whorl, some considerably stronger than others. Aperture ovate to subquadrate, constricted at posterior sinus and at junction with siphonal canal. Parietal wall thin, smooth, concave, with 1I-4 weak, oblique folds near anterior end of columella and rather weak node at edge of posterior sinus, folds and nodes sometimes absent on immature shells. Outer lip thin, broadly arcuate, slightly crenulated by termini of spiral cords, internal wall with about 12-16 lirae, those toward posterior side generally smooth and entire, those toward anterior side often periodically constrict- ed or interrupted as dashes and dots; smooth area separating tips of lirae from edge of outer lip. Anterior end of aperture constricted by anteriormost columellar plica and prominent node on inside of outer labral wall in mature shells. Siphonal canal of mature shells moderately long, slightly curved and canted to left in apertural view, smooth within; parietal margin distinct, slightly raised, forming narrow pseudoumbilicus near tip; 12-14 thin, oblique spiral cords crossed by numerous fine axial growth increments continuing from base of body whorl to tip. Operculum of lectotype ovo-elongate, subreniform, dimensions 10.6 X 5.5 mm, corneous, brown, with terminal nucleus; outer surface covered with densely packed microscopic growth lines; inner surface smooth, with large, ovate muscle scar surrounded by thick callus, about 6 distinct concentric growth increments evenly spaced across surface. Radula: see Figure 3, after Bullock (1968: 107, pl. 8, fig. 7). Distribution: Western Atlantic Ocean; southern Carib- bean Sea at Barbados, Colombia, Panama, and Nicaragua; depth range 73—220 m. Remarks: Shells of Lamellilatirus ceramidus are readily separable from all other western Atlantic Peristerniinae by the combination of characters stated in the generic diagnosis. However, L. ceramidus shells show an interesting resemblance to several species of Fusolatirus from the Indo-west Pacific, especially F. elsiae (Kil- burn, 1975) of southeastern Africa. Subsutural lamellae on shells of L. ceramidus are much more prominent than those on shells of F. e/siae, and the radula of F. elsiae (see Snyder and Bouchet, 2006:2, fig. 1-L) is Page 228 The Veliger, Vol. 50, No. 3 Figure 1. Lamellilatirus ceramidus (Dall, 1889a), lectotype, 46.2 mm, Barbados, 13°11’ 54” N, 59°38'45” W, depth 134 m, USNM 87069. Figure 2. Lamellilatirus ceramidus (Dall, 1889a), 51.0 mm, in front of Bellair Research Institute, St. James coast, Barbados, depth 220 m, ANSP 416323. Figure 3a. Lamellilatirus ceramidus (Dall, 1889a), radula (after Bullock, 1968:pl. 8 fig. 7). Figure 3b,c = Fusinus colus (Linnaeus, 1758), radula (after Barnard, 1959:fig. 19j); (b) is from an immature specimen and (c) is froma mature specimen. W.G. Lyons & M. A. Snyder, 2007 decidedly Peristernia-like, whereas the radula of L. ceramidus (Figure 3) is not. The illustration of a specimen (USNM 87069) as the holotype of Fusinus ceramidus (Dall, 1889a) by Abbott and Dance (1982; 1986) constituted a valid designation of lectotype according to International Code of Zoological Nomenclature Article 74b then in effect (ICZN, 1985), so a similar but later designation by Hadorn and Rogers (2000) was redundant. Dall (1890) chose this same shell to illustrate the species, and Bullock (1968) figured it as the holotype. We did not examine the two paralectotypes (MCZ 7240) that were examined by Hadorn and Rogers (2000), but we did examine the 7.2-mm syntype (MCZ 7239) that those authors identified as a probable juvenile of an unidentified Fusinus species. We concur that the shell is not conspecific with Fusus ceramidus, but we are not confident that it can be assigned to Fusinus or even to Fasciolariidae. Most valid records of Lamellilatirus ceramidus have involved the population at Barbados, but Hadorn and Rogers (2000) also reported three specimens trawled off Panama, Nicaragua, and the Colombian Basin; pho- tographs of the latter two specimens provided to us by Hadorn confirm their identities as L. ceramidus. Records of this species at Bermuda (Snyder, 1984, 2003; Hadorn and Rogers, 2000; Mallard and Robin, 2005) are incorrect and involve one or two species that we describe as new below. Lightbournus gen. nov. Type species: Lightbournus russjenseni, sp. nov., Recent, Bermuda, designated herein. Diagnosis: Peristerniine gastropods with small fusiform shells (lengths to 35.7 mm) and smooth, paucispiral protoconchs; whorls sculpted with broad, strong axial ribs and less prominent spiral cords; sutures distinct, not bordered by spiral threads or axial lamellae; aperture ovate, constricted anteriorly and posteriorly; parietal shield of mature shells with distinct posterior node, columella with several low but distinct, oblique folds near anterior end; inside of outer lip with well- developed lirae that are paired and entire in posterior — Page 229 half of aperture, unpaired and somewhat interrupted in anterior half; siphonal canal slender, moderately extended, canted to left in apertural view; operculum and radula unknown. Etymology: The genus name is masculine and honors J. R. H. “Jack” Lightbourn of Hamilton, Bermuda, whose ardent pursuit of Bermuda mollusks is com- memorated by this and several other taxa. Remarks: Lightbournus, known only by its type species, is readily distinguished from Lamellilatirus, with which it has been confused, by its lack of subsutural lamellae; although species of both genera have oblique columel- lar plicae, those of Lamellilatirus are broader, lower, and much less distinctly defined than those of Light- bournus. The anterior location and low, oblique shape of the columellar plicae distinguish species of Light- bournus from species of Hemipolygona, Polygona, and Pustulatirus, other western Atlantic Latirus-like genera whose columellar plicae are generally stronger, aligned more nearly perpendicular, and situated higher on the columella (see Figure 19). Lightbournus russjenseni sp. nov. Figures 4-5 Fusinus ceramidus: Snyder, 1984: 28, 30; Hadorn & Rogers, 2000: 14 (in part); Mallard & Robin, 2005: pl. 18, fig.; [zon Lamellilatirus ceramidus (Dall, 1889a)]. Fusus ceramidus: Snyder, 2003: 64 (in part); [non Lamellilatirus ceramidus (Dall, 1889a)]. Type material: Holotype, 35.7 mm, south shore of Bermuda, depth 200-240 m, traps, ANSP 416321. Paratypes: 5 sh, 33.2, 30.2, 29.9, 27.8 & 25.0 mm, 0.8 km southeast of St. Davids, south shore of Bermuda, depth 293-366 m, USNM 819198; 1 sh, 28.2 mm, 2.4 km south of Gurnet Rock, south shore of Bermuda, depth 200-240 m, traps, ANSP 416375; 2 sh, 30.3 & 26.2 mm, same locality and depth, ANSP 416374; 1 sh, 26.9 mm, same data, KLS; 1 sh, 20.7 mm, south shore of Bermuda, depth 180-240 m, traps, ANSP 416376; 3 sh, 30.3, 26.4 & 25.7 mm, 4.0 km off Figure 4. Lightbournus russjenseni sp. nov., holotype, 35.7 mm, in traps off south shore of Bermuda, depth 220-240 m, ANSP 416321. Figure 5. Lightbournus russjenseni sp. nov., paratype, 33.1 mm, in traps, Bermuda, depth 220 m, BMSM 15070. Figure 6. Bullockus mcmurrayi (Clench & Aguayo, 1941), 42.9 mm, off West End, Grand Bahama, depth 214 m, WGL. Figure 7. Bullockus mcemurrayi (Clench & Aguayo, 1941), 60.0 mm, off West End, Grand Bahama, depth 402-420 m, BMSM 26305. Figure 8. Bullockus guesti, sp. nov., holotype, 28.6 mm, trapped with hermit crab off south shore of Bermuda, depth 220 m, FLMNH 41161. Figure9. Bullockus guesti, sp. nov., paratype, 27.2 mm, in traps 2.4 km south of Gurnet Rock, south shore of Bermuda, depth 200— 250 m, ANSP 416322. Figure 10. Bullockus guesti, sp. nov., 29.7 mm, dived south of Tucker’s Town, Bermuda, depth 51 m, USNM 1100736. Page 230 The Veliger, Vol. 50, No. 3 south shore of Bermuda, depth 220 m, DMNH 096984; 2 sh, 28.8 & 25.7 mm, same data, DMNH 187105; Ish, 29.0 mm, same data, DMNH 096993; 2 sh, 28.6 & 26.1 mm, same data, DMNH 202581; 1 sh, 26.5 mm, offshore of Bermuda, traps, DMNH 212752; 1 sh, 34.4 mm, Bermuda, depth 220 m, traps, HGL; 1 sh, 33.1 mm, same data, BMSM 15070; 1 sh, 32.0 mm, same data, FLMNH 41160; 2 sh, 33.0 & 29.4 mm, Bermuda, depth 220 m, traps, WGL. Type locality: Off south shore of Bermuda, depth 200— 240 m. Description: Adult shell of moderate size, length to 35.7 mm, uniformly white, with nearly ten rapidly expanding, well-separated convex whorls. Protoconch relatively slender, of about 2-1/4 to 2-1/2 elevated whorls, first two whorls glassy, smooth, final 1/4 to 1/2 whorl with 3 or 4 rather broad axial riblets, junction with teleoconch abrupt. Nearly 8 teleoconch whorls each bearing 7, occasionally 8, broad, well-developed axial ribs crossed by less conspicuous spiral cords. Axial ribs extending from anterior suture nearly to posterior suture, slightly shouldered posteriorly, more strongly developed toward anterior suture, increasing in size anteriorly, continuing undiminished over body whorl. Suture well-defined, undulating slightly in accord with adjacent ribs and intercostal areas, rarely with faint crenulations caused by growth increments on posterior edge of adjacent anterior whorl. Spiral cords smooth, beginning at junction with protoconch; first teleoconch whorl with 4 cords, weaker 2 on posterior slope, stronger 2 crossing axial ribs, more swollen on ribs than in intercostal areas; number of cords increasing by intercalation anteriorly, about 3-4 weak ones on slope and 6 stronger ones on ribs of penultimate whorl; about 12-14 cords of more or less even strength on body whorl, continuing anteriorly onto siphonal process. Aperture ovate-elongate, constricted near posterior sinus and at intersection with siphonal canal. Parietal wall concave, smooth, with 1-3 low, oblique folds (plicae) near anterior end of columella and _ single prominent node at posterior sinus. Outer lip arcuate, finely crenulate in accord with termini of spiral cords, internal wall with 10-12 prominent to weak lirae, those toward posterior half of aperture often paired, entire, those toward anterior side on mature shells often interrupted into dashes or raised dots; lirae terminating in swollen tips before reaching edge of outer lip of mature shells, interval between tips and edge smooth. Constriction at anterior end of aperture formed by anteriormost columellar fold and prominent node on labral wall of larger shells. Siphonal canal well developed, moderately long, canted to left in apertural perspective, smooth within, with distinct parietal margin and 10-12 thin, oblique spiral cords on outer surface, cords diminishing in strength toward tip. Operculum and radula unknown. Etymology: The species name honors the late Russell H. “Russ” Jensen (1918-2001), former Emeritus Head of the Mollusk Department of the Delaware Museum of Natural History and a specialist on the Mollusca of Bermuda. Distribution: Western Atlantic Ocean; known only at Bermuda, depth range 180-366 m. Remarks: We did not ascertain how the USNM paratypes of Lightbournus russjenseni were collected, but all other specimens we examined were shells brought by hermit crabs into traps set in deep water as described by Lightbourn (1991). Snyder’s (1984) report of Fusinus ceramidus in depths of 183—366 m was based on shells of L. russjenseni, including some listed among our material examined and was the basis for Snyder’s (2003) inclusion of Bermuda in the range of Fusus ceramidus. More recently, Mallard and Robin (2005) published two color photographs of a Bermudan shell of L. russjenseni that they identified as Fusinus ceramidus. Hadorn and Rogers (2000) also included Bermuda within the range of F. ceramidus, but without indicating the source of their information; their record was probably of L. russjenseni. Lightbournus russjenseni may also have been represented among shells that Lightbourn (1991) listed as Latirus brevicaudatus; we examined a few L. russjenseni among shells received (by MAS) from Lightbourn as L. brevicaudatus, but most of those shells represent another new species described later in this paper. Bullockus gen. nov. Type species: Bullockus guesti sp. nov., Recent, Bermuda, designated herein. Diagnosis: Peristerniine gastropods with broadly fusi- form shells of small to large size for subfamily (lengths 30-82 mm); whorls sculpted with moderate to strong axial ribs and less prominent spiral cords; sutures incised, lacking adjacent cords or lamellae; siphonal canals relatively short, slender, canted to left, with shallow pseudoumbilicus near tip; apertures ovate, constricted anteriorly and posteriorly, with parietal shields with columellar plicae absent, rudimentary, or developed only on largest mature shells; and outer lips of mature shells bearing internal lirae interrupted as dashes or dots. Etymology: The genus name is masculine and honors Dr. Robert C. Bullock, University of Rhode Island, Kingston, RI, whose earlier studies of western Atlantic Latirus-like taxa paved the way for our study. W. G. Lyons & M. A. Snyder, 2007 Remarks: Species of Bullockus differ from other Latirus-group species by the combination of characters defined in the diagnosis. Species other than the type species that we classify in Bullockus include Latirus (Hemipolygona) mcmurrayi Clench and Aguayo, 1941, from northern Cuba and the northwestern Bahama Islands, depths 214-420 m (Clench and Aguayo, 1941; Lan, 1993: Sunderland and Sunderland, 1996; Petuch, 2002; Snyder, 2006; this report); Hemipolygona honkeri Snyder, 2006, from the southwestern Caribbean Sea and the eastern Bahamas, depths 245—550 m (Snyder, 2006); Latirus (Latirus) varai Bullock, 1970, from northeastern Cuba, depth 183 m (Bullock, 1974); and Bullockus pseudovarai sp. nov., from the eastern Bahama Islands, depths 245—550 m. All of these species share features of the columella and suture that characterize Bullockus. The type species previously was mistaken for a species of Polygona Schumacher, 1817, but shells of that genus have prominent, near-perpendicular colu- mellar plicae and sutures bordered with spiral cords, wrinkles, axial lamellae, or often all three. The other included species previously were classified with Latirus Montfort, 1810, and Hemipolygona Rovereto, 1899, but are distinguished from Latirus by having shells with angulate whorls bearing distinct spiral cords and subovate apertures and from Hemipolygona by lacking distinct columellar plicae and by having sutures that are finely incised between the smooth surfaces of adjacent whorls, without adjacent cords, wrinkles or lamellae. Latirus mcmurrayi differs from other species of Bullockus by having a shell with an often remarkably expanded umbilicus. This is a variable feature in several latirid genera. Similar aberrations are found in some specimens of Hemipolygona recurvirostris (Schubert & Wagner, 1829). Some shells of L. mcmurrayi have a few weak, oblique folds or even small, tooth-like plicae near the anterior end of the columella, but on most shells the columella is as featureless as that of the type species of Bullockus; that feature and the complete lack of ornamentation around the suture prompt us to place the species in Bullockus. Vermeij and Snyder (2006: 417) had tentatively placed this species in Hemipoly- gona. We observed several kinds of variation among eleven specimens of B. mcmurrayi that we examined: holotype, 52.2 mm, off Matanzas, Cuba, 348 m, MCZ 135285; 1 sh, 73.5 mm, off Matanzas, Cuba, 400-420 m, KLS; 1 sh, 55.7 mm, off Tamarind, Grand Bahama Island, ANSP 368995; 2 sh, 52.3 & 39.7 mm, off Tamarind, Grand Bahama, 214 m, KLS; | sh, 42.9 mm, off West End, Grand Bahama, 214 m, WGL; | sh, 54.5 mm, off West End, 408-421 m, WGL (Figure 6); 1 sh, 41.5 mm, off West End, 26°38’N, 78°59'W, 420 m, ANSP 416377; and 3 sh, 48.6 mm (BMSM 26298), Page 231 39.0 mm (BMSM 26299) & 60.0 mm (BMSM 26305; Figure 7), all off West End, Grand Bahama, 402— 420m. The holotype and KLS shells from off Matanzas, Cuba, have weak spiral cords, causing them to appear smoother than most Bahamian shells, but cords on the WGL and two of the three BMSM shells from off West End are nearly as weak as the Cuban specimens. The columella of the holotype lacks any indication of plicae, but the other shell from off Matanzas, by far the largest specimen examined at 73.5 mm, has three low, tooth-like plicae at the anterior end of the columella; one to three rather vague columellar folds are present on the nine shells from Grand Bahama. Finally, the holotype is darkly stained (by mud?) and the large KLS shell from off Matanzas is grayish-white, but most shells from Grand Bahama are uniformly light yellow; the exceptions, two shells from off West End (BMSM 26299 & 26305; Figure 7) are covered with horizontal brown bands caused by the presence of that color over all areas not occupied by white spiral cords. Features of Hemipolygona honkeri that prompt us to reclassify the species in Bullockus are discussed in remarks for B. guesti. Similarly, features of Latirus varai that place it in Bullockus are discussed with the account for B. pseudovarai. Bullockus guesti sp. nov. Figures 8—10 [?] Latirus sp. near sanguifluus ““Rve.” Peile, 1926: 82; Bullock, 1974: 76; [non Latirus sanguifluus (Reeve, 1847), Recent, Polynesia, = Turrilatirus sanguifluus, fide Vermeij & Snyder, (2006: 419)]. Latirus brevicaudatus (Reeve, 1847). Waller, 1973: 43; Lightbourn, 1991: 5; [non Latirus brevicaudatus (Reeve, 1847), = Polygona brevicaudata (Reeve, 1847), fide Vermeij and Snyder (2006: 420)]. Type material: Holotype 28.6 x 12.8 mm, off south shore of Bermuda, depth 220 m, trapped with hermit crab, FLMNH 41161. Paratypes: 29.7 13 mm, south of Tucker’s Town, Bermuda, depth 51 m, dived by T. Waller, USNM 1100736; 1 sh, 27.2 mm, 2.4 km south of Gurnet Rock, south shore of Bermuda, depth 200— 250 m, traps, ANSP 416322; 4 sh, 25.1, 23.1, 18.7 & 16.2 mm, south shore of Bermuda, depth 180-240 m, traps, ANSP 416373; 1 sh, 20.3 mm, same data, BMSM 15071; 1 sh, 19.3 mm, same data, KLS; 1 sh, 22.0 mm, same locality, 200-240 m, traps, ANSP 416372; 1 sh, 19.4mm, 4.0 km off south shore of Bermuda, depth 220 m, traps, DMNH 234001; 2 sh, 20.3 & 18.5 mm, south of Gurnet Rock, Bermuda, depth 220 m, DMNH 187106; 1 sh, 21.2 mm, Ber- muda, depth 220 m, traps, WGL. Page 232 Description: Shell solid, broadly fusiform, to 29.7 mm long, 13.0 mm wide, with about 10 whorls. Protoconch smooth, glassy, of about 2 whorls, with rounded tip and convex sides; final 1/2 whorl with 3-5 axial riblets of increasing strength; junction with teleoconch abrupt. Teleoconch pale orange, sometimes faded nearly white, of about 8 rapidly expanding, sharply angled whorls, with prominent axial ribs, less prominent orange- brown spiral cords, and a well-marked suture. Axial ribs about 8 per whorl, broad, abutting anterior suture and extending to, but not over, posterior sutural ramp. First teleoconch whorl with about 3 spiral cords of equal size: cords increasing by intercalation to 6 or more on second and later whorls, including about 4 smaller cords on sutural ramp and posterior halves of ribs and 2 larger cords on anterior halves of ribs, sometimes with another cord adjacent to anterior suture; 3-4 largest cords on each whorl colored dark orange-brown, contrasting with lighter background color of teleoconch; body whorl with as many as 12 dark cords, 2 cords at periphery strongest. Suture well- marked, undulating in accord with ribs and intercostal areas, without subsutural lamellae. Aperture ovate to subquadrate; parietal wall smooth, concave, distinctly demarked on mature shells, with node-like callus at posterior sinus; columella without plicae but terminating in (usually) sharp angle at junction with siphonal canal. Outer lip arcuate, finely crenulate in accord with termini of spiral cords, internal wall with 8—10 well-developed lirae, posterior lirae entire, anterior ones interrupted as dashes or dots on mature shells, lirae not extending to lip edge; node at anterior end usually prominent, uncommonly reduced, together with columellar terminal angle forming constriction between aperture and siphonal canal. Siphonal canal rather short, slender, canted to left in apertural view, smooth within; edge of parietal callus distinct, raised on larger shells, forming shallow, chink- like pseudoumbilicus near anterior tip; about 7 dark or lighter smooth oblique cords continuing from base of body whorl to tip. Operculum (of paratype USNM 11000736) ovate, brown, corneous, dimensions 5.4 X 3.6 mm, nucleus terminal, with concentric growth increments on upper surface and muscle scar bordered by conspicuous callus beneath, callus wider and thicker near nucleus. Radula unknown. Distribution: Western Atlantic Ocean: known only at Bermuda, depth range 51—250 m. Etymology: The species name honors the late Arthur Tucker Guest, O. B. E. (1907-1993), retired customs officer and student extraordinaire of the shells of Bermuda, who was instrumental in collecting and distributing most shells that we examined. The Veliger, Vol. 50, No. 3 Remarks: Except for the USNM paratype collected at a depth of 51 m using scuba (Waller, 1973:fig. 11), all specimens we examined of Bullockus guesti were occupied by hermit crabs taken in traps set in deeper water as described by Lightbourn (1991). The depth of collection of Waller’s specimen is much shallower than any other recorded for the species and is the only depth where a living specimen has been collected. Shells of B. guesti have been confused with Polygona brevicaudata (Reeve, 1847). Although their shells are similar in general profile, in sculpture of the axial ribs, and particularly in having orange-brown spiral cords, P. brevicaudata can be distinguished by having distinct, well-developed, near-perpendicular plicae on its colu- mella and rugose wrinkles, often overlain by fine axial lamellae, just anterior to its suture. The only reports of Latirus living at Bermuda involve Peile’s (1926) uncertain listing of “‘Latirus sp. near sanguifluus (Rve.),” Waller’s (1973) listing of Latirus brevicaudatus from a depth of 51m off the southern coast, and Lightbourn’s (1991) mention of crabbed shells of L. brevicaudatus among specimens trapped in deep waters off the southern coast. Bullock (1974) proposed that Peile’s record may have been based on Latirus angulatus (Réding, 1798), in which he included L. brevicaudatus as a junior synonym. “Latirus” sanguifluus (Reeve, 1847), reclassified as Turrilatirus sanguifluus by Vermeij and Snyder (2006), is a Polynesian species endemic to the Tuamotu and Marquesas Archipelagos (Salvat and Rives, 1975) and is unlikely to occur at Bermuda. Bullock (1974) also proposed that Waller’s record of L. brevicaudatus represented L. angulatus, but we re-examined that specimen (USNM 1100736), which is B. guesti. The “L. brevicaudatus” of Lightbourn is also B. guesti, as evidenced by shells received from Lightbourn and Guest, and all specimens except Waller’s that we examined originated from that source. Thus, it seems likely that all reports of Recent P. brevicaudata at Bermuda were actually of B. guesti. Reports of L. brevicaudatus in Bermuda Pleistocene deposits are errors for a species of Leucozonia (see discussion). Shells of Bullockus guesti most resemble those of B. honkeri, (Figure 14), but shells of the latter species are uniformly yellow and larger (to 55.5 mm, versus 29.7 mm for B. guesti), with larger protoconchs, whorls with more prominent spiral cords, colored the same as other exterior surfaces, that protrude and create the appearance of points where they cross axial ribs, and several inconspicuous folds on the columella. Bullockus pseudovarai sp. nov. Figures 11,13 Latirus varai: Snyder, 2006: 41; [non Latirus (Latirus) varai Bullock, 1970, Recent, northeastern Cuba]. W. G. Lyons & M. A. Snyder, 2007 Page 233 Figure 11. Bullockus pseudovarai, sp. nov., holotype, 82.3 mm, off San Salvador, Bahama Islands, depth 488 m, ANSP 416379. Figure 12. Bullockus varai (Bullock, 1970), possible paratype, 57.2 mm, off Gibara, Oriente Province, Cuba, depth 183 m, KLS. Figure 13. Bullockus pseudovarai, sp. nov., paratype, 64.7 mm, off San Salvador, Bahama Islands, depth 220-550 m, ANSP 416378. Figure 14. Bullockus honkeri (Snyder, 2006), holotype, 39.6 mm, off San Salvador, Bahama Islands, depth 245-550 m, ANSP 413204. Page 234 The Veliger, Vol. 50, No. 3 Hemipolygona varat: Snyder, 2006: 44, pl. 1, figs. la—d, 2a—d: [non Latirus (Latirus) varai Bullock, 1970]. Type material: Holotype 82.3 < 27.7 mm, from off San Salvador, Bahama Islands, depth 488 m, ANSP 416379; paratype 64.7 mm, off San Salvador, Baha- mas, depth 220-550 m, ANSP 416378. Type locality: Off San Salvador, eastern Bahama Islands, 488 m. Description: Shell solid, relatively large (to 82.3 x 27.7 mm) fusiform, with about 11 whorls. Protoconch of about 2 smooth, glassy whorls, with rounded tip and convex sides, about 4 axial riblets on final 1/2 whorl; junction with teleoconch abrupt. Teleoconch white with prominent orange-brown axial ribs and other patches of similar color scattered on body whorl; about 9 postnuclear whorls increasing in size anteriorly; axial ribs well developed, 7 ribs on early whorls, increasing to 8 by about whorl 5; ribs aligned in transverse rows between whorls beginning at about whorl 5. Spiral cords 3 on first whorl, of about equal size, anterior- most 2 cords stronger than others thereafter, creating “‘squared”’ effect where they cross ribs and conferring tabulate appearance to spire; number of cords increas- ing to 4 on about whorl 5, with some barely perceptible secondary threads between, middle 2 cords strongest, resembling points where they cross ribs; about 11 cords of varying strength on body whorl, 2 cords at shoulder much stronger than others. Suture incised, distinct, undulating slightly in accord with adjacent ribs and interspaces, bordered above and below by essentially smooth shell surfaces interrupted only by very fine, irregularly spaced axial growth increments. Aperture ovate, white, with slight anterior and posterior constrictions; parietal wall essentially smooth, concave, slightly thicker at posterior sinus; columella bearing single, inconspicuous, oblique fold, arching anteriorly in gradual transition to siphonal process. Outer lip arcuate, marked with 2 points near shoulder and lesser crenulations elsewhere in accord with termini of spiral cords of body whorl; internal wall with as many as 20 lirae of unequal size, some terminating before others, none extending to edge of lip, termini of farthest extending lirae punctuated with extra dots at end; prominent node at anterior edge of liral array, forming constriction at entrance to siphonal process. Siphonal canal rather short, slender, canted to left in apertural view, smooth within; inner edge (continua- tion of parietal wall) distinct, raised, forming shallow, slender pseudoumbilicus near anterior tip; about 8 white, oblique cords continuing from base of body whorl to tip, diminishing in strength anteriorly. Operculum brown, corneous, fairly slender, curved and tapering anteriorly to terminal nucleus; outer surface marked with numerous closely packed, arc-like concentric growth increments; inner surface with muscle scar bearing concentric elliptical rings of growth, surrounded by thick ring of callus. Radula unknown. Etymology: The name pseudovarai is formed of the Greek prefix pseudes, meaning false, and varai, the name of a similar species; literally, the false varai, referring to a report by Snyder (2006) in which the new species was mistakenly figured and discussed as Hemipolygona varai. Distribution: Western Atlantic Ocean; known only from near San Salvador, Bahama Islands, depths 220-550 m. Remarks: Snyder (2006: 44, pl. 1, figs. 1, 2) figured two specimens of Bullockus pseudovarai as Hemipolygona varai, a Closely related species from northeastern Cuba. The smaller of the specimens (64.7 mm) that Snyder figured is the paratype of the new species; the larger of the shells (75.2 mm) is in the collection of Tom Honker of Delray Beach, Florida. The only variations noted among those two shells and the holotype is that the anterior-most of the two shoulder cords on the 75.2- mm shell is somewhat weaker than the posterior cord, conferring to the whorl a less “‘squared” profile; the axial ribs are less aligned with each other on consecutive whorls of the 75.2-mm shell than on the other two. Snyder (2007) shows that the name Latirus varai was incorrectly applied by Pointier & Lamy (1998: 131) and Mallard & Robin (2005: pl. 51) to shells from the Lesser Antilles, and Snyder has reclassified that species in the genus Hemipolygona. A shell from Venezuela that Mallard & Robin (2005: pl. 51) also figured as Latirus varai appears to be undescribed. We believe that the only valid reports of Latirus varai involve the 70 mm holotype and a paratype from off Gibara, Oriente Province, Cuba, in 183m. The holotype, MCZ 262589, has been illustrated by Bullock (1970: text-fig. 1), Abbott (1974: text-fig. 2493a), Kaicher (1978: card 1838), Abbott & Dance (1982, 1986: 186, row 1, right fig.), Petuch (1987: pl. 8, fig. 13), Snyder (2000: fig. 3), and Vermeij & Snyder (2006: 418, fig. 3A). The paratype, which remained in the collection of John Finlay of Wilmington, Delaware, has not been figured. A specimen of Latirus varai from off Gibara in 183 m that was acquired from Finlay by the Sunder- lands was illustrated by Sunderland & Sunderland (1996: 17). We expected that shell to be the sole paratype, but its size is 57.2 mm, not 52.4 mm as reported for the paratype by Bullock. Whatever its status may be, the Sunderland shell is clearly conspe- cific with the holotype and we reillustrate it here (Figure 12) for comparison with B. pseudovarai. W. G. Lyons & M. A. Snyder, 2007 Bullockus varai can be distinguished from B. pseudovarai by having many more white spiral cords crossing the brown axial ribs; the Sunderland shell has 5 cords crossing ribs on teleoconch whorls 3 and 4, 7 cords on whorls 5 and 6, 9 cords on whorl 7, and 12-13 cords on the body whorl. The cords are smaller and more closely spaced than those of B. pseudovarai and, because of their subequal size, do not create the “squared” profile seen on B. pseudovarai. Instead, the ribs of B. varai are rather evenly convex. We concur with Bullock’s (1970) observation that the general appearance of the ribs of B. varai is much like that of Latirus kandai Kuroda, 1950 (= Fusolatirus kandai; Recent, Japan and Philippine Islands), but ribs of B. pseudovarai certainly do not resemble those of F. kandai. Bullock (1970: 134) described four folds, “‘the upper one weaker,” on the holotype of B. varai, and two of those folds are evident in the original figure. However, folds of B. varai are weaker, more oblique, and more anteriorly situated than are the near-perpendicular folds (plicae) of Hemipolygona and Polygona species. The Sunderland shell has no folds on its columella, evidently because it is less mature. General shell shape, including those of the protoconch, the columella, the featureless shell surfaces around the sutures, and the form of the siphonal process all dictate placement of Latirus varai in Bullockus. DISCUSSION All of the species treated herein have been associated at some time with Latirus Montfort, 1810, a generic name that has served as an umbrella for a diverse array of taxa distributed throughout tropical and subtropical regions of the world. As consequences of recent revisions noted in the introduction, many and in fact most of those species are now classified elsewhere, leaving relatively few Indo-west Pacific species and no western Atlantic species in Latirus (see Vermeij and Snyder, 2006). New World species until recently classified in Latirus are now placed in Polygona, Hemipolygona, and Pustulatirus, genera whose shells, together with those of Leucozonia and Opeatostoma Berry, 1958, have columellas bearing three or more well-developed, near-perpendicular plicae (Figures 15— 18). Prominent columellar plicae also occur on shells of all species of the Indo-west Pacific genera Turrilatirus, Latirolagena Harris, 1897, and Peristernia Morch, 1852, and on most species of Latirus as construed by Vermeij and Snyder (2006). These plicae are prominent even on shells not yet mature. Such plicae do not occur on species of the Indo-west Pacific genera Benimakia or Fusolatirus, nor do they occur on the new genera Lamellilatirus and Lightbournus. Plicae may occur, uncommonly, on some mature shells of Bullockus varai, Page 235 and B. mcmurrayi, but most shells of that species lack plicae, and no plicae have been seen on mature shells of B. guesti, B. honkeri, or B. pseudovarai. The marine molluscan fauna of Bermuda clearly is derived from the Caribbean region (Jones, 1876), but its remote location and cool winter climate seem to have acted as impediments to recruitment by many species. Groups such as the Fasciolariidae, with direct development or only brief planktotrophic larval stages, have shown very limited success recruiting to Bermuda. Although many species of Fasciolariidae are known from the Caribbean Sea and Gulf of Mexico, only three (Fusinus lightbourni Snyder, 1984; Lightbournus russ- Jenseni, sp. nov.; and Bullockus guesti, sp.nov.) are known to live at Bermuda today, all in fairly deep water (~50—250 m) and all evidently endemic to Bermuda. Although we have identified no close relative of L. russjenseni, the nearest relative of F. lightbourni seems to be F. schrammi (Crosse, 1865) from deep water in the northeastern Antilles, and B. guesti belongs to a group that includes several species that live in deep waters (183-550 m) of the Bahama Islands, Cuba and the southwestern Caribbean Sea (Snyder, 2006). If there are no shallow-water fasciolariids living at Bermuda today, historical records and reports of Pleistocene fossils suggest that a few such species reached that remote island, located ~1000 km to the east of North America (Muhs ef al, 2002). Shallow- water Caribbean species reported from Bermuda include Fasciolaria distans Lamarck, 1816 (= Fascio- laria liliwum Fischer von Waldheim, 1807), reported by Jones (1864, 1876); Fasciolaria tulipa (Linnaeus, 1758), reported by Moore and Moore (1946) and Richards er al. (1969); Leucozonia nassa (Gmelin, 1791) and its junior synonym L. cingulifera (Lamarck, 1822), report- ed by Heilprin (1889), Peile (1926), and Moore and Moore (1946); and Latirus brevicaudatus (Reeve, 1847), reported by Richards et al. (1969) and Waller (1973). Dall (1889b) tentatively listed Bermuda within the range of Leucozonia ocellata (Gmelin, 1791), but that listing has not been substantiated. There are also two unfortunate Bermuda listings of the Indo-west Pacific Pleuroploca trapezium (Linnaeus, 1758) by Santos Galindo (1977: 190, 372) but those listings are obviously spurius and may be ignored. Fasciolaria tulipa and Leucozonia nassa are the two most widely ranging western Atlantic fasciolariids; F. tulipa occurs now from northeastern Brazil to North Carolina, and L. nassa ranges from Trindade Island, 1140 km off the Brazilian coast (Vermeij and Snyder, 2002) and throughout central and northern Brazil to North Carolina (Nathanson, 2006). Despite limitations imposed by their modes of reproduction and dispersal, these species have demonstrated abilities to recruit over distances that seem to constitute barriers for most other Page 236 The Veliger, Vol. 50, No. 3 SE Figure 15. Figure 16. Figure 17. Figure 18. Figure 19. 416321. 18 ‘4 19 Pustulatirus mediamericanus (Hertlein & Strong, 1951), 78.0 mm, off Acapulco, Mexico, ANSP 416383. Hemipolygona armata (A. Adams, 1855), 47.7 mm, Goree Island, Dakar, Senegal, depth 15-30 m, ANSP 416384. Polygona infundibulum (Gmelin, 1791), 70.7 mm, Cabo de la Vela, Guajira, Columbia, depth 60 m, ANSP 416385. Bullockus memurrayi (Clench & Aguayo, 1941) 55.6 mm, Tamarind, Grand Bahama Island, depth 245 m, ANSP 368995. Lightbournus russjenseni, sp. nov., holotype, 35.7 mm, in traps off south shore of Bermuda, depth 200-240 m, ANSP W. G. Lyons & M. A. Snyder, 2007 fasciolariids, so their presence in Bermuda may seem reasonable. However, although reports indicate that both species may have arrived at Bermuda from time to time, each has encountered difficulty establishing a population there. Fasciolaria was first reported at Bermuda by Jones (1864), who cited F. distans as rare based on one specimen “‘in a semifossil state.”” Jones (1876) reiterated that record and added that his marine specimens had been identified by the well known conchologist C. B. Adams, although Adams died in 1853 (Abbott, 1973), more than a decade before Jones published his first list. The Jones record was repeated by Dall (1885; 1889a, b), Heilprin (1889), and Verrill (1907). Peile (1926) also repeated the fossil record of F. distans and added that the species still lived at Bermuda but was rarely found; Peile’s was the last report of F. distans at Bermuda. The next record of Fasciolaria, by Moore and Moore (1946), reported F. tulipa as occurring both fossil and living at Bermuda, although the new records were implicitly of fossil shells which had been identified by W. J. Clench. Moore and Moore noted the shells to be fairly common in subsurface calcareous sandstone cut by harbor dredging from below sea level in and around Castle Harbour, the age of the sandstone being considered ‘‘preglacial” or possibly a result of rising sea level within a glacial period. Abbott (1958) later treated the report of F. distans by Peile (1926), and implicitly others, as F. tulipa and repeated earlier comments that living specimens were rare at Bermuda. Finally, Richards ez al. (1969) reported new records of F. tulipa in the Pleistocene Belmont Formation (age ~200 ka; Muhs et a/., 2002) of Bermuda but noted that the species is not living in Bermuda waters today, an observation reinforced by Sterrer (1998). Thus, it seems to have been decided that the original report of F. distans as a Bermuda fossil was an error for F. tulipa, that the sole evidence of living Fasciolaria at Bermuda was Peile’s unsubstantiated note, and that no species of Fasciolaria now lives at Bermuda. We have not seen any museum specimens of Recent Fasciolaria at Bermuda, but we did examine a voucher (ANSP 59434) from Watch Hill Park, Bermuda, that Richards et al. (1969) reported as a Quaternary fossil of F. tulipa. The specimen is an upper fragment of a spire, height 25.0 mm, with its early whorls intact. The specimen is not fasciolariid but may be a fragment of a species of Cassidae, perhaps Casmaria. This finding refutes the only record of Fasciolaria at Bermuda known to be supported by physical evidence and provokes uncertainty about other Bermuda records of the genus. Richards et al (1969) also reported Bermuda Pleistocene records of Latirus brevicaudatus at Spen- cer’s Point (Spencer’s Point Formation, age 130,000 + 15,000 ybp) and Grape Bay (Devonshire Formation; = Page 237 Devonshire marine member of Rocky Bay Formation, age ~125 ka; Muhs et a/., 2002), and they noted that a report by Moore and Moore (1946) of fossil Lewcozonia cingulifera Lamarck in Castle Harbour dredgings may instead have been L. brevicaudatus. Based on the report by Richards et al, Muhs et al. (2002: 1372) cited Latirus brevicaudatus as one of three extralimital southern (warm-water) gastropod species that ranged northward to Bermuda during the last interglacial period but do not live around Bermuda today. We examined specimens representing both of the Richards et a/. records, now catalogued as Latirus sp. at the Academy of Natural Sciences of Philadelphia. The voucher from Spencer’s Point (ANSP 61785) is a small shell fragment consisting of most of a body whorl and including the aperture and columella. Prominent features include a few large, nodose ribs on the shoulder crossed by 2 fairly prominent spiral cords, followed anteriorly by 5—6 much lower cords and | markedly larger cord near the anterior edge of the whorl. There are about 7 finer spiral threads on the subsutural ramp but no indication of subsutural lamellae. The columella has 3, possibly 4 near- perpendicular plicae. The voucher from Grape Bay (ANSP 61768) is a larger (36.3 21.2 mm), nearly intact shell missing only the anterior end of the siphonal process but encased in limestone concretions over much of the outer surface and aperture. There are about 8 node-like ribs per whorl; several small, narrow cords on the subsutural ramp are overlain by micro- scopic growth increments, and the columella is obscured by concretion. We compared both vouchers with Recent Leucozonia nassa in the ANSP collection and judge them to be conspecific. These fossils constitute the only physical evidence we have seen of the presence of Leucozonia nassa at Bermuda. Heilprin (1889: 168) first reported L. nassa (as L. cingulifera) at Bermuda without comment, indicating that he did not consider the occurrence noteworthy, and Peile (1926) also listed L. cingulifera among Bermudan mollusks collected by him and Arthur Haycock. Moore and Moore (1946) mentioned L. cingulifera as rare among living fauna of Bermuda and also reported shells from two “‘preglacial”’ [Pleis- tocene] beds there. But then Richards eft al. (1969) suggested that Pleistocene records by Moore and Moore were actually of L. brevicaudatus, casting doubt on other previous records, none of which can now be substantiated. No more reports of L. nassa at Bermuda since those by Moore and Moore have been forthcom- ing, but our findings support those earlier reports and refute the Pleistocene records of Latirus brevicaudatus. Having also refuted one fossil record of Fasciolaria tulipa by Richards et al. and knowing of no other specimens of that species to demonstrate its Bermuda occurrence, we conclude that L. nassa may be the only Page 238 shallow-water Caribbean fasciolariid species to have recruited to Bermuda since before the onset of the last ice age. Acknowledgments. Robert C. Bullock, University of Rhode Island, granted permission to reproduce the figure of the radula of Fusus ceramidus from his unpublished thesis and provided other information from that study to the authors. Jack Lightbourn of Hamilton, Bermuda, shared information about his deep-water trapping program. Roland Hadorn of Lyss, Switzerland, provided photographs of specimens in his collection. Dr. Timothy A. Pearce, Carnegie Museum of Natural History, provided information on Russell Jensen, on the molluscan collection at DMNH, and on Bermudan molluscan literature. Liz Shea and Leslie Skibinski (DMNH), Alex Baldinger (MCZ), and Paul Greenhall, Warren Blow, and M. G. “Jerry” Harasewych (USNM) are thanked for locating specimens and facilitating loans of material from their institutions. Harry G. Lee of Jacksonville, Florida, and Kevan and Linda Sunderland of Sunrise, Florida, loaned specimens for study from their personal collections. 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THE VELIGER a B The Veliger 50(3):241-247 (October 1, 2008) © CMS, Inc., 2007 Injuries on Nautilus Jaws: Implications for the Function of Ammonite Aptychi ISABELLE KRUTA' AND NEIL H. LANDMAN? "Muséum National d’Histoire Naturelle, Paris, France; Université Pierre et Marie Curie, Paris VI, CNRS-UMR 5341 Paléobiodiversité et paléoenvironnements, Case 104-4, Place Jussieu, 75252, Paris Cedex 05, France ? Division of Paleontology (Invertebrates), American Museum of Natural History, 79" St. and Central Park West, New York, New York 10024 (e-mail: landman@amnh.org) Abstract. Documentation of repaired injuries and abnormalities on the jaws of modern nautilus sheds light on the ecology and behavior of these animals. It also helps elucidate the function of ammonite aptychi, which are traditionally interpreted as opercula. We examined 219 pairs of jaws belonging to Nautilus belauensis, N. macromphalus, N. pompilius, and Allonautilus scrobiculatus. Abnormalities occur in 68% of the sample and are only present on the lower jaw. The abnormalities consist of 1) repaired fractures, 2) small depressions, 3) radial grooves and ridges, and 4) flexures in the chitin. These abnormalities are either the result of injury or growth pathology. Injuries may be due to accidents during feeding (e.g., biting down on a hard crustacean carapace) or from predatory attacks. Alternatively, they may have been sustained during mating behavior or fighting between males. Most abnormalities occur on the left side of the lower jaw. This may be related to the fact that in male nautilus, the jaws are displaced to the right side of the midline, so that during mating, for example, the apex of the jaws of the male lines up with the left side of the jaws of the female. The presence of injuries and other abnormalities on the jaws of nautilus suggest that similar features on aptychi may have been produced during the normal use of the jaws, and do not necessarily imply an opercular function. Alternatively, aptychi may have served to strengthen and reinforce the lower jaw. INTRODUCTION Aptychi are present in many Jurassic and Cretaceous ammonites, mainly the Ammonitina and Ancylocer- atina (Lehmann, 1980; Engeser and Keupp, 2002; Tanabe and Landman, 2002; Landman et al., 2006). Aptychi are pairs of calcitic plates that cover the outside surface of the outer lamella of the lower jaw (Figure 1A). Traditionally, aptychi have been inter- preted as opercula (Trauth, 1927-1936; Schindewolf, 1958; Seilacher, 1993; Keupp, 2007), but the discovery of aptychi with other elements of the buccal mass demonstrated their homology with the lower jaws of present-day cephalopods (Lehmann, 1975, 1980). Nevertheless, the opercular theory has never been completely discarded, and as a compromise, Lehmann and Kulicki (1990) have suggested a double function, with aptychi serving as both jaws and _ opercula. According to these authors, the aptychus was capable of rotating into a nearly vertical position to act as an operculum in the event of an attack by a predator. The main pieces of evidence for the interpretation of aptychi as opercula are (1) the close match between the size and shape of the aptychus and the aperture of the ammonite, 2) the calcitic composition and ornamenta- tion of the apytchus, suggesting a protective function, and (3) the presence of repaired injuries on the aptychus. Such injuries have been interpreted by Engeser and Keupp (2002) as resulting from predatory attacks. This assertion implies that injuries are only present on opercula, not jaws. The protective function of opercula in gastropods is well known (see, for example, Vermeij and Williams, 2007, and references therein). In order to evaluate the significance of repaired injuries on aptychi, we studied the externally shelled cephalopod nautilus to determine if repaired injuries or other abnormalities are present on the jaws of these animals. The function of the jaws in nautilus is known. They serve for biting and tearing food. An opercular function is performed instead by the fleshy hood, which is composed of thick connective tissue. The presence of repaired injuries on the jaws of nautilus would imply that injuries, by themselves, are not sufficient proof of an opercular function. Such injuries could equally result from trauma to the jaws during the lifetime of the animal. The presence of injuries on nautilus jaws also provides insights into the ecology of these reclusive organisms. Because of the deep water habitat of nautilus, direct observations of their behavior are problematic. As a result, investigators have relied on indirect evidence, including analyses of the isotopic The Veliger, Vol. 50, No. 3 upper jaw rhyncholite aptychus il lower jaw conchorhynch lower jaw Ammonite Jaw Nautilus Jaw Figure 1. Comparison of the jaws of ammonites with those of modern nautilus. A. Reconstruction of the upper and lower jaws of the Early Jurassic ammonite Hildoceras (after Lehmann, 1975: fig. 4). Note that the lower jaw is covered with a pair of calcareous plates, known as an aptychus. il = inner lamella; ol = outer lamella. B. Upper and lower jaws of nautilus (after Tanabe and Fukada, 1999: fig. 19.3). The tips of the jaws are reinforced with calcareous deposits. 11 = inner lamella; ol = outer lamella. - composition of the shell (e.g., Cochran et al., 1981) and studies of repaired injuries (e.g., Arnold, 1985), for clues about their life history and habitat, MATERIAL AND METHODS The upper and lower jaws of nautilus are composed of chitin and each consists of inner and outer lamellae (Figure 1B). The upper jaw sits inside the lower jaw. The oral opening is surrounded by the labial margin (‘lips’). The outside surfaces of the outer lamellae of the upper and lower jaws are mostly covered by a thin membrane of connective tissue and epithelium (Tanabe and Fukuda, 1999). In contrast, the inside surface of the outer lamella and the outside surface of the inner lamella of the lower and upper jaws are covered with muscles. The jaws are reinforced with calcareous deposits at the apex, known as rhyncolites and conchorhynchs (for a more complete discussion of the orientation and terminology of nautilus jaws, see Saunders et al., 1978). We studied 219 pairs of nautilus jaws in the collec- tions of the American Museum of Natural History (AMNH). All of the jaws are presumably from adult specimens. Part of the sample (164 specimens) consisted of the entire buccal mass preserved in alcohol. The other 55 pairs of jaws were already separated from the buccal mass and subsequently dried. In the alcohol preserved specimens, the upper jaws were still covered by tissue and were not examined for marks. However, in the dried specimens, both the upper and lower jaws were inspected. The jaws belong to four species of Nautilus: 141 jaws of Nautilus macromphalus from New Caledonia (57 males, 44 females, 40 indet.); 34 jaws of Nautilus belauensis from Palau; 42 jaws of Nautilus pompilius from various localities (3 from Samoa, 4 from Indonesia, 3 from Fiji, 24 from Papua New Guinea, 8 from the Philippines); and 2 jaws of Allonautilus scrobiculatus from Papua New Guinea. In addition to repaired breaks, we recorded the presence of other abnormalities on the jaws. Specimens were analyzed under the stereomicroscope (x6—x50) and six specimens were observed with SEM. Because of the loss of flexibility of the chitin after drying, some parts of the jaws were broken. These breaks were easy to recognize because of the freshness and sharpness of the fractures. RESULTS Description of Abnormalities We categorized the abnormalities observed on the nautilus jaws as follows: (1) repaired fractures, (2) depressions, (3) radial grooves and ridges, and (4) flexures. A total of 149 specimens (68%) of the sample exhibit at least one of these abnormalities. Almost all of the abnormalities occur on the outer lamella of the lower jaw. No abnormalities are present on the upper jaw. 1) Repaired fractures: The most spectacular repaired fracture appears on a lower jaw of Nautilus belauensis (AMNH _ 51881). The jaw is 39mm long and is undoubtedly from an adult specimen. The injury occurs on the left side of the outer lamella and extends from the anterior dorsal part of the wing to the posterior ventral part (Figure 2A, B). The break continues onto the inner lamella (Figure 2C). The outside of the outer lamella is fractured, with a maximum offset of 1.3 mm on each side of the break. On the inside of the outer lamella, the fracture is repaired by an elongate, cordlike thickening of chitin (Figure 2C, D). This chitinous thickening is 2.2mm wide and 1.0mm high and subdivides for part of its length. There are no growth lines on the thickening and the texture is reminiscent of pahoehoe lava. 2) Depressions: Small depressions are present on some of the nautilus jaws (Figure 3C—F, I, J). These features only occur on the outside of the outer lamella of the lower jaw. Several depressions may occur on the same specimen (Figure 3E, F). The depressions are less than 1 mm long and less than 0.2 mm deep. They vary in shape from triangular to quadrate to round. They are conformable with the surrounding jaw surface, but show a slightly different texture. The depressions are not expressed on the inside surface of the outer lamella. They occur in all of the nautilus species but are more common in Nautilus pompilius and N. belauensis. They preferentially occur on the left side of the outer lamella in these species. I. Kruta & N. H. Landman, 2007 Page 243 Figure 2. Lower jaw of Nautilus belauensis (AMNH 51881) with a large repaired injury. A. Left lateral view of the lower jaw, showing the break (arrow) on the outer lamella (ol). The break extends to the posterior end of the wing. Part of the calcareous deposit at the tip was broken away during handling. Anterior direction to the left. B. Left lateral view of the lower jaw, slightly tilted down, to expose the gap at the break (arrow). C. View of the inside surface of the outer lamella, showing the repaired portion (left arrow). Note that the break (right arrow) continues onto the inner lamella (il). Anterior direction to the right. D. Close-up of the repair (arrow) on the inside surface of the outer lamella, which consists of a thickened ridge of chitin that must have been secreted from the inside. 3) Radial grooves and ridges: As noted by Saunders et al. (1978), the outside surface of the outer lamella of the lower jaw is covered with closely spaced, comarginal lirae, which are usually interpreted as growth lines. In some specimens, however, these growth lines are transected by elongate grooves that occasionally extend to the posterior margin (Figure 3A, B). These grooves are generally superficial with a maximum depth of approximately 0.2 mm, and are usually bordered by thin ridges. Sometimes, instead of grooves, the surface is marked by thin bands, approximately 0.5 mm wide, characterized by an irregular texture (Figure 3G, H). Radial grooves or bands occur in 54% of the specimens that show abnormalities. 4) Flexures: Flexures are minor discontinuities in the outer lamella of the lower jaw, which follow the course of the lrae (Figure 3K, L). They are sometimes expressed as overhanging fringes of chitin, indicating the previous position of the jaw margin. Flexures are very common and preferentially occur on the left side of the lower jaw (92% of the specimens with flexures). Page 244 The Veliger, Vol. 50, No. 3 Figure 3. Abnormalities on the lower jaws of nautilus. A, B. Radial groove (arrow) on the outer lamella of the lower jaw of Nautilus macromphalus (AMNH 51870). The groove extends to the posterior margin of the wing, suggesting permanent damage to the jaw-secreting tissue. Anterior direction to the right. The close-up in B is rotated 180° relative to A. C, D. Elongate depression (arrow) on the outer lamella of the lower jaw of Nautilus belauensis (AMNH 51868). Anterior direction to the left. E, F. Triangular depression (arrow) on the outer lamella of the lower jaw of Nautilus belauensis (AMNH 51335). Anterior direction to the right. G, H. Radial bands (arrows) on the outer lamella of the lower jaw of Nautilus belauensis (AMNH 51883). Anterior direction to the right. I, J. Small depression (arrow) on the outer lamella of the lower jaw of Nautilus pompilius (AMNH 51869). Anterior direction to the left. The close-up in B is rotated 180° relative to I. K, L. Flexure in the chitin (arrow) on the outer lamella of the lower jaw of Nautilus macromphalus (AMNH 51884). Anterior direction to the right. The calcareous tips of the jaws are occasionally missing in these specimens due to breakage during drying or handling. Distribution of Abnormalities The incidence of abnormalities varies among the different species. The highest percentage of abnormal- ities occurs in Nautilus belauensis, including the specimen with the conspicuous scar. The highest percentage of grooves appears in N. macromphalus. The percentage of depressions is higher in N. be/auensis and N. pompilius than in the other species. In N. macromphalus, in which the number of males and females is known, the incidence of abnormalities is higher in males than in females (70% versus 55%). The most common abnormality in both sexes is grooves. However, the incidence of grooves is higher in males than females (60% versus 36%). Some of these differences may be related to the preservation of the jaws (alcohol versus dry). For example, most of the flexures occur in alcohol- preserved rather than dry specimens (92% versus 8%), perhaps because flexures are less noticeable on dry specimens. In contrast, depressions are more common on dry specimens. Therefore, the kind of preservation may bias the results. DISCUSSION The abnormalities described on the jaws of nautilus are either the result of injuries to the jaws or growth pathologies. The repaired break in AMNH 51881 is the most obvious example of a repaired injury in which the lower jaw was broken and repaired during life by the secretion of additional chitinous material from the inside. The depressions, with a maximum depth of 0.2 mm, may also represent injuries due to impact, or I. Kruta & N. H. Landman, 2007 Page 245 possibly damage caused by parasites. In any event, the damage was not permanent. In contrast, the radial grooves and ridges extending to the posterior margin of the jaws imply permanent damage to the jaw-secreting tissue, perhaps due to injury. The flexures that follow the growth lines probably represent pauses in growth due to stress, after which growth resumed, and are not related to injuries to the jaws. The injuries on these jaws may be due to several factors. They may have been produced during feeding. In their natural habitat, nautilus regularly feed on crustaceans (Saunders and Ward, 1987; Saunders et al., 1987; Ward, 1987), as confirmed by reports of crusta- cean remains in the gut contents of freshly captured nautilus (Haven, 1972). In addition, Ward and Wicksten (1980) observed Nautilus macromphalus in captivity eating freshly molted exoskeletons of lobsters. Thus, the nautilus may have damaged their jaws by simply biting down on a hard crustacean carapace. Injuries may also have been caused by counterattacks of the prey. For example, Ward (1981) observed hermit crabs defending themselves against nautilus attack by breaking off pieces of the nautilus shell. Alternatively, the injuries on nautilus jaws may have been produced by predators. The large breaks observed on nautilus shells have routinely been attributed to predators such as teleosts, sharks, and crabs (Arnold, 1985; Ward, 1987). There are few eye witness accounts of predatory attacks, but Saunders et al. (1987) observed such an attack by triggerfish on Nautilus belauensis in shallow water in Palau. Another possible source of injuries on nautilus jaws may be linked to mating and courtship behavior. During copulation, a male nautilus grasps the shell of the female (Arnold, 1985, 1987), and the jaws of both sexes could be damaged in the process. The jaws of the male are especially vulnerable to counter attack by the female if the male is using its jaws to grasp the shell of the female. Injuries could also be produced during fights between males. Haven (1972) noted bites in the hoods and V-shaped breaks in the shells of Nautilus pompilius in captivity, which she attributed to combat between males. This behavior is consistent with the observation that in our sample of N. macromphalus, in which the distribution of sexes is known, jaw abnor- malities are more common in males than females. The location of the abnormalities sheds some light on the biology of nautilus. Nearly all of the abnormal- ities occur on the outer lamella of the lower jaw. Because the upper jaw is mostly covered by muscles and sits within the lower jaw, the outer lamella of the lower jaw is more vulnerable to injury. Furthermore, most of the abnormalities occur on the left side of the lower jaw. This pattern may be related to the position of the jaws in the shell. In males, the jaws are displaced toward the right side of the shell due to the development of the spadix (Saunders and Spinosa, 1978; Saunders and Ward, 1987). Thus, during copulation, the apex of the jaws of the male lines up with the left side of the jaws of the female and, conversely, the apex of the jaws of the female lines up with the left side of the jaws of the male. This offset has also been cited to explain the disparity in the incidence of repaired shell breaks between the left and right sides of the shell. For example, Arnold (1985) noted that, in females, there is a higher incidence of injuries on the left side of the shell. The off-center position of the jaws in males also implies that, during male to male combat, the left side of the jaws is more vulnerable to damage than the right side. The highest percentage of abnormalities occurs in Nautilus belauensis. This probably reflects the larger size of the jaws of this species. With more surface area to inspect, the probability of finding injuries is higher. In addition, studies of the longevity of nautilus suggest that NV. belauensis is longer lived than the other nautilus species (Landman and Cochran, 1987). Because of their longer life span, these animals may have had a greater chance of sustaining injuries. The paleontological implications of our observations bear on the arguments used to support the opercular theory of ammonite aptychi (Seilacher, 1993; Keupp, 2007). Traditionally, irregular marks on aptychi have been interpreted as healed injuries and, thus, proof of an opercular function. There are many illustrations of such marks on aptychi from the Jurassic of Germany (for example, Schindewolf, 1958: pls. 5, 9; Keupp et al., 1999: pl. 3, fig. 6; Keupp, 2000: 114, upper left; Engeser and Keupp, 2001: fig. 2). In addition, Landman et al. (2007: figs. 13.17—20) have illustrated such marks on the aptychi of Baculites from the Upper Cretaceous of North America. Our study of abnormalities on nautilus jaws suggests that the formation of such features on aptychi may have been the result of the normal use of the jaws. However, we cannot exclude the possibility of an opercular function, although this interpretation re- quires more evidence. Alternatively, the aptychi may have simply served to strengthen the lower jaw. In this context, it 1s worth noting that nearly all of the abnormalities that we observed on nautilus jaws appear on the outside surface of the outer lamella of the lower jaw. Thus, the thick calcareous plates comprising the aptychus may have functioned to protect the outer surface of the lower jaw in ammonites, even if the aptychus did not rotate into a vertical position, as envisioned by Lehmann and Kulicki (1990). FUTURE WORK This is the first study of abnormalities on nautilus jaws (or any cephalopod jaws, for that matter). Further Page 246 studies could investigate the relationship between injuries on the nautilus shell and those on the jaws. Is an injury on the shell also expressed on the jaws? Such studies might provide additional insights into the ecology and behavior of nautilus—that is, their prey and their predators. It would also be interesting to determine if there are geographic differences in the occurrence of jaw injuries associated with different feeding habits. From the paleontological point of view, our study suggests the need to more carefully examine the abnormalities on ammonite aptychi. Are these marks, in fact, the same as those on nautilus jaws? Do they appear on the inner and outer sides of the aptychus or only on the outer side? Do the injuries extend to the underlying chitinous layer of the jaw or are they restricted to the calcareous plates? The extent to which the marks on aptychi are the same as those on the jaws of nautilus will determine the extent to which our observations about nautilus can shed light on the functional interpretation of ammonite jaws. Acknowledgments. We thank Royal H. Mapes (Ohio Univer- sity) and W. Bruce Saunders (Bryn Mawr College) for supplying specimens for our study. R.H. Mapes, Gary Vermeij (University of California, Davis), Kazushige Tanabe (Univer- sity of Tokyo) and Isabelle Rouget and Fabrizio Cecca (both Université Pierre et Marie Curie, Paris) reviewed an earlier draft of this manuscript and made many helpful suggestions. Jacob Mey (AMNH) assisted in SEM, Susan Klofak (AMNH) in specimen preparation, and Jay Biederman and Steve Thurston (both AMNH) in photography. This research was funded by the Norman D. Newell Fund (AMNH) and a fellowship to I. Kruta from the Kade Foundation to support her stay at the AMNH. LITERATURE CITED ARNOLD, J. M. 1985. Shell growth, trauma, and repair as an indicator of life history for Nautilus. The Veliger 27(4): 386-396. ARNOLD, J. M. 1987. Reproduction and embryology of Nautilus. Pp. 353-372 in W. B. Saunders & N. H. Landman (eds.), Nautilus: the biology and_ paleobi- ology of a living fossil. Plenum Press: New York and London. COCHRAN, J. K., D. M. RYE & N. H. LANDMAN. 1981. Growth rate and habitat of Nautilus pompilius inferred from radioactive and stable isotope studies. Paleobiology 7:469-480. ENGESER, T. & H. Keupp. 2002. Phylogeny of the aptychi- possessing Neoammonoidea (Aptychophora nov., Ceph- alopoda). Lethaia 24:79—96. HAVEN, N. 1972. The ecology and behavior of Nautilus pompilius in the Philippines. The Veliger 15(2):75-81. Keupp, H. 2000. Ammoniten: Palaobiologische Erfolgsspir- alen. Jan Thorbecke Verlag: Stuttgart. 165 pp. Keupp, H. 2007. Complete ammonoid jaw apparatuses from the Solnhofen plattenkalks: implications for aptychi function and microphagous feeding of ammonoids. 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Jaws of Late Cretaceous placenticeratid ammonites: How preser- vation affects the interpretation of morphology. American Museum Novitates 3500:1-48. LEHMANN, U. 1975. Uber Nahrung und Ernahrungsweise von Ammoniten. Palaontologische Zeitschrift 49:187— 195. LEHMANN, U. 1980. Ammonite jaw apparatus and soft parts. Pp. 275-287 in M. R. House & J. R. Senior (eds.), The Ammonoidea. Academic Press: London. LEHMANN, U. & C. KULICKI. 1990. Double function of aptychi (Ammonoidea) as jaw elements and opercula. Lethaia 23:325—-331. SAUNDERS, W. B. & C. SPINOSA. 1978. Sexual dimorphism in Nautilus from Palau. Paleobiology 4:349-358. SAUNDERS, W. B., C. SPINOSA & L. E. DAVIS. 1987. Predation on Nautilus. Pp. 201-212 in W. B. Saunders & N. H. Landman (eds.), Nautilus: the biology and _paleo- biology of a living fossil. Plenum Press: New York and London. SAUNDERS, W. B., C. SPINOSA, C. TEICHERT & R. C. BANKS. 1978. The jaw apparatus of Recent Nautilus and _ its palaeontological implications. Palaeontology 21(1):129— 141. SAUNDERS, W. B. & P. D. WARD. 1987. Ecology, distribution, and population characteristics in Nautilus. Pp. 137-162 in W. B. Saunders & N. H. Landman (eds.), Nautilus: the biology and paleobiology of a living fossil. Plenum Press: New York and London. SCHINDEWOLF, O. H. 1958. Uber Aptychen (Ammonoidea). Palaeontographica A 111:1-46. TANABE, K. & Y. FUKUDA. 1999. Morphology and function of cephalopod buccal mass. Pp. 245-262 in E. Savazzi (ed.), Functional morphology of the invertebrate skeleton. John Wiley & Sons: New York. TANABE, K. & N. H. LANDMAN. 2002. Morphological diversity of the jaws of Cretaceous Ammonoidea. Pp. 157-165 in H. Summesberger, K. Histon & A. Daurer (eds.), Cephalopods—Present and Past. Abhandlungen der Geologischen Bundesanstalt 57. TRAUTH, F. 1927-1936. Aptychenstudien I-VIII. Annalen des Naturhistorischen Museums in Wien 41:171—259 (1927); 42: 121-193 (1928); 44: 329-411 (1930); 45: 17-136 (1931); 47: 127-145 (1936). SEILACHER, A. 1993. Ammonite aptychi: how to transform a jaw into an operculum? American Journal of Science 293A:20-32. I. Kruta & N. H. Landman, 2007 VERMEW, G. T. & S. T. WILLIAMS. 2007. Predation and the geography of opercular thickness in turbinid gastropods. Journal of Molluscan Studies 73:67-73. WARD, P. D. 1981. Shell sculpture as a defensive adaptation in ammonoids. Paleobiology 7:96—100. Page 247 WARD, P. D. 1987. The natural history of Nautilus. Allen & Unwin: London. 267 pp. WARD, P. D. & M. K. WICKSTEN. 1980. Food sources and feeding behaviour of Nautilus macromphalus. The Veliger 23(2):119-124. THE VELIGER © CMS, Inc., 2007 The Veliger 50(3):248—254 (October 1, 2008) Effect of Temperature and Feeding Preference on Submerged Plants by the Island Apple Snail, Pomacea insularum (d’Orbigny, 1839) (Ampullariidae) L. A. GETTYS, W. T. HALLER, C. R. MUDGE AND T. J. KOSCHNICK University of Florida Department of Agronomy, Center for Aquatic and Invasive Plants, 7922 NW 71 Street, Gainesville, FL 32653. Tel: 352-846-2516; Fax: 352-392-3462 (e-mail: lgettys@ufl.edu) Abstract. The island apple snail (Pomacea insularum (d’Orbigny, 1839)) is a South American snail that became naturalized in Florida waterways in the mid-1970s and has recently spread throughout much of the state. Food consumption by this herbivorous snail was determined in 10-day feeding trials at temperatures of 15 to 35°C. Optimum feeding of the exotic submerged plant Aydrilla verticillata (L.f.) Royle (hydrilla) occurred over a wide temperature range (20 to 35°C). However, snail growth was greatest at temperatures of 20 to 30°C. Free choice plant preference studies were conducted to determine feeding preferences for native and exotic submerged plants. One exotic and two native species (17. verticillata, Najas guadalupensis (Spreng.) Magnus (southern naiad) and Chara sp. (stonewort), respectively) were highly preferred by island apple snails, followed by the two native species Potamogeton illinoensis Morong. (Illinois pondweed) and Vallisneria americana Michx. (tapegrass). Leaves of the exotic species Myriophyllum aquaticum (Vell.) Verdc. (parrotsfeather) were eaten after the more preferred plants were consumed and no significant feeding was noted on the exotic species Egeria densa Planch. (Brazilian elodea). While island apple snails have distinct preferences for certain submerged plants, they consumed both native and exotic species, which may significantly affect growth of certain species and will likely change species composition of submerged plant communities in Florida wherever they are common. Key Words: canaliculata complex, feeding preference, Florida waterways, invasive species, temperature effect. INTRODUCTION The island apple snail (Pomacea insularum (a Orbigny, 1839)) is native to South and Central America and was most likely introduced into Florida through the aquarium trade (Thompson, 1997). Collections of this exotic snail have been reported sporadically in natural areas of Florida since 1978 (FLMNH, 2007), but recent reports suggest that the species has greatly expanded its range and has invaded virtually all counties in central and southern Florida (Denson, 2005). Specimens of P. insularum have been collected from a variety of habitats in Florida, including natural and constructed wetlands, streams, ponds, lakes and ditches that are inundated for most or all of the year. These habitats have earthen bottoms and are characterized by the presence of submerged, floating and/or emergent aquatic macro- phytes. Areas frequented by P. insularum also typically have structures above the water line (e.g., emergent aquatic plants or man-made structures such as dikes or locks) onto which P. insularum deposits eggs. There has been some confusion regarding the taxonomy of South American apple snails in Florida, but recent DNA studies indicated that the snails present in Florida (originally identified as P. canalicu- lata (Lamarck, 1822)) are actually P. insuwlarum (Rawlings et al., 2007). Field identification of species within the family Pomacea is challenging, since external characters are highly plastic and influenced by envi- ronmental factors. This is especially true of P. haustrum (Reeve, 1856) (titan), P. canaliculata (channeled) and P. insularum (island) apple snails, which are virtually indistinguishable from one another in morphology and behavior. In fact, these snails are so similar they are often grouped together into a “canaliculata complex.” Cazzaniga (2002) stated that all canaliculata-like apple snails may constitute a single, highly variable species and further noted that any canaliculata-like apple snail has the potential to become a pest and cause damage to aquatic ecosystems. Canaliculata-like apple snails can become quite large, but P. insularum is considered the largest of the group and can attain a shell height of up to 150 mm (Benson, 2008; Gettys and Haller, 2007). One factor that influences the feeding of canaliculata- like apple snails is temperature. Estebenet and Cazza- niga (1992) and Estebenet and Martin (2002) found that P. canaliculata grew most quickly under warm conditions (>25°C) and stopped feeding during cool temperatures (e.g., <18°C). These results suggest that the warm year-round temperatures found throughout much of Florida may provide an ideal habitat for P. insularum; therefore, the first objective of this experi- L. A. Gettys et al., 2007 ment was to measure the effect of temperature on consumption of Aydrilla_ verticillata (L.f.) Royle (hydrilla), an aquatic weed that is ubiquitous in aquatic systems throughout Florida, Texas and other regions that have been invaded by P. insularum. Canaliculata-like snails in the genus Pomacea are voracious herbivores and have been introduced to some parts of the world as biocontrol measures to manage aquatic weeds (Cowie, 2002; Okuma et al., 1994; Wada, 1997). Some workers have reported that canaliculata- like apple snails actively select their food and exhibit a preference for some plant material. For example, Fukushima et al. (2001) found that P. canaliculata preferred most fruits and vegetables to rice seedlings. Lach et al. (2000) stated that P. canaliculata selected Vigna marina (Burm.) Merr. (beach pea or notched cowpea) over Ejichhornia crassipes (Mart.) Solms (waterhyacinth), Ludwigia octovalvis (Jacq.) P.H. Ra- ven. (primrose willow) and Pistia stratiotes L. (water- lettuce). Carlsson and Lacoursiere (2005) stated that P. canaliculata virtually eliminated Lemna minor L. (duckweed) and E. crassipes after 6 and 21 days of grazing, respectively, but reduced the biomass of Ipomea aquatica Forsk. (waterspinach) by only 20% after 32 days of grazing. Estebenet (1995) found that P. canaliculata preferred Zannichellia palustris L. (horned pondweed) over Myriophyllum elatinoides Gaudin (water milfoil or Christmas-tree plant) and Chara contraria A. Braun ex Kutz. (opposite stonewort); snails had a low preference for Rorippa nasturtium- aquaticum [L.] Hayek (watercress) and Potamogeton striatus Ruiz & Pavon (broadleaf pondweed) and did not select Elodea canadensis LC Rich. in Michx (common waterweed). In contrast, Cazzaniga and Estebenet (1984) and Cowie (2002) suggested that these snails feed indiscriminately on virtually anything, including algae, macrophytes, phytoplankton, detritus and even immature snails of other species. In addition, Peltzer and Lajmanovich (2003) reported that Hyla pulchella Dumeéril & Bibron, 1841 (anuran tadpoles) were consumed by juvenile P. canaliculata. A variety of submerged macrophytes species are found in Florida; some species are native, while others are exotic and invasive. Common native aquatic macrophytes in Florida include Vallisneria americana Michx. (tape- grass), Najas guadalupensis (Spreng.) Magnus (south- ern naiad), Potamogeton illinoensis Morong. (Illinois pondweed), and Chara sp. (stonewort). Invasive exotic species common in Florida waters include H. verticil- lata (native to Asia, Africa and Australia), along with the South American natives Egeria densa Planch. (Brazilian elodea) and Myriophyllum aquaticum (Vell.) Verdc. (parrotsfeather). Native species of macrophytes are more desirable than exotic species, but many aquatic fauna that rely on submerged vegetation as a habitat for nesting and spawning do not discriminate Page 249 between native and exotic species. If the canaliculata- like P. insularum indiscriminately consumes all flora, the consequences to Florida’s ecosystem could be devastating since the snail can cause significant changes to wetland ecosystems through herbivory of aquatic macrophytes. High densities of P. canaliculata in natural wetlands in Thailand caused an almost complete loss of aquatic macrophytes through grazing (Carlsson et al., 2004). Wetlands in central and south Florida persist under the same environmental condi- tions as those in Thailand, so it is reasonable to expect the same situation to occur if waterways in Florida become infested with P. insularum. Data regarding the feeding habits and macrophyte preferences of canali- culata-like apple snails are conflicting and there are no reports that address the feeding habits of snails positively identified as P. insularum; therefore, the second objective of this study was to determine if P. insularum is truly a non-specific feeder or if the snail shows a feeding preference when presented with a diversity of native and exotic submerged macrophytes commonly found in Florida waterways. MATERIALS AND METHODS Several hundred specimens of P. insularum were collected from a heavily infested earthen irrigation pond (surface area 0.12 ha; maximum depth | m) at a wholesale aquatic plant nursery in Lake City, Florida. Snails were maintained in a covered greenhouse (ambient temperature 28 + 3°C) at the University of Florida’s Center for Aquatic and Invasive Plants in Gainesville, FL for 2 to 3 weeks prior to their utilization in these experiments. Each snail was measured (height and width), weighed and assigned a letter/number combination code using the following system. Snails were assigned one of six letter classes based on weight (A: <25 g; B: 25 to 35 g; C: 35.1 to 45 g; D: 45.1 to 55 g; E; 55.1 to 65 g; and F: >75 g) and were numbered in ascending order (e.g., snail D6 weighed between 45.1 and 55 g and was the 6™ snail labeled in weight class D). The shell of each snail was gently cleaned and dried using a disposable paper towel, then the alphanumeric code was painted onto the snail shell using nail polish so that snails could easily be identified. This coding system allowed positive identification of each snail and was used to ensure that each snail was only used in a single study. Effect of Temperature on Consumption The objective of this study was to measure the effect of temperature on consumption of macrophytes by P. insularum. The macrophyte H. verticillata was used in this study because it is an exotic species that is ubiquitous in aquatic systems throughout Florida, Page 250 Texas and other regions that have been invaded by P. insularum. The temperature regimes in this study were chosen to represent the range of seasonal variation in water temperature in Florida. Consumption of H. verticillata under five temperature regimes (15°C, 20°C, 25°C, 30°C and 35°C) was investigated in growth chambers (Percival model E36L, Perry, IA). Digital controls on the chambers were programmed to maintain a daylength of 14 hr and to hold water temperature at the target temperature + 0.5°C. Four 5- gallon aquaria were placed in each growth chamber. Water was maintained at a depth of ca. 25 cm to provide a water volume of ca. 12 L and aeration was supplied by a standard aquarium aerator. Snails were selected for uniform weight and one snail was placed in each filled aquarium within the growth chamber. Water temperature was adjusted by 3°C per day to reduce shock during the transition from ambient greenhouse temperature (28 + 3°C) to the assigned experimental temperature regime. Snails were acclimatized to exper- imental temperatures for 2, 4 or 6 d for 25°C and 30°C treatments, 20°C and 35°C treatments and 15°C treatment, respectively, and were fed H. verticillata ad libitum during acclimatization. Snails were starved for 24 hr after acclimatization and were weighed prior to commencement of each study. Each study lasted 10 d and each snail was offered a total of 90g of H. verticillata during each study (30 g each on days 1, 4 and 7). Water in the aquaria quickly became fouled with feces and detritus, so additional aquaria were filled with water and placed in each growth chamber on days 3 and 6. Snails, aerators and uneaten H. verticillata were moved to these clean, acclimatized tanks on days 4 and 7. Total biomass consumption was calculated by weighing uneaten plant material remain- ing in each aquarium on day 10 of each study and snails were weighed on day 10 as well. Data were analyzed to detect differences in plant biomass consumption and differences in snail weight under each temperature regime. Feeding Preference Macrophytes were maintained in a greenhouse under natural daylength during Fall 2005 at the University of Florida’s Center for Aquatic and Invasive Plants in Gainesville, FL. All macrophytes were grown in square pots (10 cm square X 12 cm deep) filled with 1 kg of coarse builder’s sand amended with 1 g of Osmocote® Plus 15-8-12 controlled-release fertilizer (The Scotts Co. LLC, Marysville OH). Seven submerged macrophytes — FH. verticillata, V. americana, E. densa, N. guadalupensis, P. illinoensis, Chara sp. and M. aquaticum — were utilized in this experiment to represent some of the most common native and exotic aquatic species found in Florida waters. Macrophytes were propagated by The Veliger, Vol. 50, No. 3 vegetative means with either four 10-cm-long apical cuttings per pot (1. verticillata, E. densa, N. guadalu- pensis, P. illinoensis and M. aquaticum), one clump of ten 10-cm-long apical cuttings per pot (Chara sp.) or three rooted plantlets per pot (V. americana). These propagation methods were used because our prelimi- nary studies suggested these protocols would supply snails with sufficient amounts of each macrophyte species. Macrophytes were propagated 7 to 14 d prior to commencement of the experiment and were grown in circular fiberglass tanks (inside diameter 105 cm, water depth 55 cm) filled with well water to a volume of ca. 475 L (pH 8, temperature range ca. 22°C to 32°C). All food was withheld from snails for 48 hr prior to commencement of the food preference experiments. Naylor (1996) stated that a density of 8 snails/m? (32,376 snails/acre) could reduce rice yields by 90%, and snail densities in our experiment bracket that of Naylor (12 snails/m* = 46,729/acre in Study 1 and 7 snails/m? = 28,037/acre in Study 2). Snails were sorted by weight and then randomly allocated to each tank so that all treatment tanks had similar amounts of snail biomass (i.e., mean weights of 658.7 + 9.2 g and 407.7 + 7.1 gin Studies | and 2, respectively). Shell height of snails used in these experiments ranged from 58 to 74 mm, and biomass per snail ranged from 50 to 86 g. Eight circular fiberglass tanks (inside diameter 105 cm, water depth 55cm) were used in this experiment with 4 pots of each macrophyte species placed in a completely randomized design in each tank. Four tanks were used as controls and contained only macrophytes, while the remaining four tanks were populated with macrophytes and P. insularum. All tanks were covered with fiberglass insect screening to ensure containment of the snails in treatment tanks and to maintain consistent light conditions between snail tanks and control tanks. Feeding preference data were collected every other day (Study 1) or every third day (Study 2). One pot of each macrophyte species was randomly removed from each tank at each data collection interval, resulting in four replicates of each treatment (control vs. snails). This allowed us to account for macrophyte growth during the course of the each study. Two replicates of this experiment were conducted in Fall 2005. Study 1 ran from 22 Sept. to 30 Sept. and Study 2 ran from 24 Oct. to 2 Nov. Experimental parameters were similar in both studies except for snail density (10 snails per tank in Study 1 and 6 snails per tank in Study 2) and macrophyte removal interval (2, 4, 6 and 8 d in Study | and 3, 6, 9 and 12 d in Study 2). Snails consumed macrophytes more quickly than anticipated in Study 1, so snail density and macrophyte removal interval for Study 2 were modified in an attempt to more clearly identify the snails’ preference among the macrophytes offered. Separate analyses were performed for each study to L. A. Gettys et al., 2007 120 100 80 60 40 Fresh weight (g) of hydrilla eaten in 10 days 15C 20C 25C 30 C 35C fe) Figure 1. Amount of H. verticillata eaten by P. insularum under various temperature regimes. Each bar represents the mean of four replicates (snails) per temperature regime and temperature regimes coded with the same letter indicates that H. verticillata consumption was not significantly different at P = 0.05. account for seasonal differences in plant growth. Each study was treated as a factorial (2 7 < 4) design, with 2 treatments (snails vs. control), 7 macrophyte species and 4 macrophyte removal intervals. Macrophyte shoot material was weighed to determine fresh weights for all treatments and percent macrophyte material eaten by snails was then calculated by comparing uneaten macrophyte weight of each species in snail tanks to the mean of the same macrophyte species in control tanks. Direct consumption data were not analyzed because macrophyte biomass varied by species (e.g., three rooted plantlets of V. americana weigh considerably more than ten 10-cm-long apical cuttings of Chara sp.), so the use of percentage data allowed us to standardize consumption across macro- phytes of disparate biomass. These percentage data were subjected to analyses of variance (SAS Version 9.1, SAS Institute Inc., Cary, NC, USA) to detect differences between treated and control plants of the same species and differences among plant species harvested at a given time interval. RESULTS Effect of Temperature on Herbivory Snails consumed less plant material at 15°C than at intermediate and high temperatures (20°C, 25°C, 30°C and 35°C) (Figure 1). Also, snails grew faster at intermediate temperatures (20°C, 25°C and 30°C) than at low and high temperatures (15°C and 35°C) (Figure 2). Snails held at 30°C consumed an average of 63.8 g of H. verticillata over the course of the study (a daily average of 10.0 g of H. verticillata per kg of snail weight) (Figure 1) and gained an average of 2.3 g of biomass over the course of the 10-day study period Page 251 44 Change in snail biomass (g) in 10 days -2 Figure 2. Change in biomass of P. insu/arum under various temperature regimes. Each bar represents the mean of four replicates (snails) per temperature regime and temperature regimes coded with the same letter are not significantly different at P = 0.05. (Figure 2). In contrast, snails kept at 15°C ate an average of 14.5 g of H. verticillata during the course of the study (a daily average of 2.3 g of H. verticillata per kg of snail biomass) (Figure 1) and actually lost an average of 1.8 g of body weight during the 10-day study period (Figure 2). These results indicate that optimum consumption of the submerged plant JH. verticillata by P. insularum occurred over a wide temperature range (20 to 35°C). However, snail growth was greatest at temperatures of 20 to 30°C. Feeding Preference Snail weight did not change significantly during the course of Study 1, as final mean biomass per tank was 655.8 + 5.7 g. Snails in Study 1 preferred N. guadalupensis, Chara sp. and H. verticillata to P. illinoensis, V. americana, M. aquaticum and E. densa; with no difference noted among WN. guadalupensis, Chara sp. and H. verticillata at any sampling interval (Figure 3a). Snails ate 89.5%, 82.4% and 75.2% of N. guadalupensis, Chara sp. and H. verticillata, respective- ly, within two days of commencement of the study and had consumed 100% of these species by the fourth day of Study 1 (Figure 3a). Snails selected P. i/linoensis and V. americana over M. aquaticum and E. densa when most-preferred macrophytes were depleted. Snails consumed M. aquaticum when most-preferred and less-preferred macrophyte species were depleted, but typically fed on leaves and not stem material. Fresh biomass of E. densa in snail tanks was not different from that in control tanks at the conclusion of the study; therefore, E. densa was not eaten by snails, even when all other plant material had been consumed. Most-preferred macrophytes in Study 1 were N. guadalupensis, Chara sp. and H. verticillata. Less- preferred macrophytes P. i/linoensis and V. americana [R= F vertioitata $B V. amaricana —e=E. densa HEN. quadalypensis —&=P. iinoansis —@= Chara sp. —t=M. aquaticum Percent macrophyte biomass consumed 2 days 4 days 6 days 8 days ==? ilinoansis —O= Chara sp. = M. aquaticum Percent macrophyte blomass consumed 3 days 6 days 9 day: 12 days Figure 3a. Percent macrophyte biomass eaten by P. insu- larum during Study |. Each point represents the mean of four replicates per species and species coded with the same letter are not significantly different at P = 0.05. Error bars indicate standard error. Figure 3b. Percent macrophyte biomass eaten by P. insu- larum during Study 2. Each point represents the mean of four replicates per species and species coded with the same letter are not significantly different at P = 0.05. Error bars indicate standard error. were consumed when most-preferred macrophytes were depleted, and least-preferred macrophytes (with little or no feeding damage) were M. aquaticum and E. densa. Snail weight did not change significantly during Study 2, as final mean biomass per tank was 410.3 + 8.0 g. Snails in Study 2 preferred N. guadalupensis and FH. verticillata to Chara sp., P. illinoensis, V. americana, M. aquaticum and E. densa, with no difference noted between N. guadalupensis and H. verticillata at any sampling interval (Figure 3b). Snails ate 81.5% and 73.2% of the N. guadalupensis and H._ verticillata, respectively, within three days of commencement of the study. Snails preferred Chara sp. to P. illinoensis, V. americana, M. aquaticum and E. densa and consumed 50.0% of Chara sp. by the third day of Study 2 The Veliger, Vol. 50, No. 3 (Figure 3b). There was no difference in consumption of N. guadalupensis, H. verticillata and Chara sp. by the sixth day of Study 2 because nearly all macrophyte material of these most-preferred species was consumed prior to day 6 (Figure 3b). Snails selected P. i/linoensis over V. americana, M. aquaticum and E. densa when most-preferred macrophytes were depleted and con- sumed V. americana in preference to M. aquaticum and E. densa when most-preferred macrophytes and P. illinoensis were scarce. As in Study 1, snails fed on leaves of M. aquaticum when preferred species were scarce and E. densa was not eaten by snails even when all other macrophyte material had been consumed. Most-preferred macrophytes in Study 2 were N. guadalupensis and H. verticillata. The less-preferred macrophyte Chara sp. was selected over P. illinoensis and V. americana, and least-preferred macrophytes (with little or no feeding damage) were M. aquaticum and E. densa. DISCUSSION This research revealed that P. insularum consumed the greatest amount of macrophyte biomass and accumu- lated the greatest amount of body weight when exposed to moderate temperatures (1.e., 20°C to 30°C). This finding does not bode well for Florida’s wetlands and waterways, since water temperature falls in this range throughout most of the year in central and southern Florida. These results also showed that P. insularum does indeed exhibit a feeding preference if offered an assortment of submerged macrophytes over a short time interval. However, it is unlikely that macrophyte origin plays a role in P. insularum’s food selection, since both native (N. guadalupensis) and exotic (H. verticil- lata) species were preferred in both studies. Three macrophyte species (including the weed H. verticillata) were most preferred, while E. densa was completely rejected by snails in this experiment, even when the majority of other macrophyte material had been consumed. It is interesting to note that H. verticillata (native to Africa or Asia and strongly preferred by P. insularum) and E. densa (native to South America and rejected by P. insularum) are both members of the Hydrocharitaceae, as is Elodea canadensis LC Rich. in Michx (common waterweed). Other researchers (e.g., Carlsson and Lacoursiere, 2005; Estebenet, 1995) have noted that P. canaliculata will not eat E. canadensis, which is native to temperate regions of North America; in fact, snails offered only E. canadensis by Estebenet (1995) refused to eat the macrophyte and eventually starved to death. These three species are submerged macrophytes that are morphologically similar and difficult to distinguish from one another, so the reason for the snails’ preference for H. verticillata and rejection of E. densa and E. canadensis is unclear. L. A. Gettys et al., 2007 It is commonly thought that macrophytes employ structural defenses (e.g., spines, thorns or toughness) to deter feeding by herbivores. Pennings and Paul (1992) found that plant toughness and calcification deterred feeding by the marine gastropod Dolabella auricularia (Lightfoot, 1786) (sea-hare). However, D. auricularia is able to sequester plant secondary metabolites that may act as chemical feeding deterrents in other herbivorous species. Chemical defenses against herbivory have been extensively studied in terrestrial plants but have only recently gained attention in aquatic macrophytes. Secondary metabolites including alkaloids, glucosi- nates, polyphenols and flavonols have been identified in a number of aquatic macrophytes and reduce or prevent consumption by herbivores. Erhard et al. (2007) found that flavonoids and other allelochemicals produced by Elodea nuttallii (Planch.) St. John (western waterweed) reduced feeding by the larvae of the generalist pyralid aquatic moth Acentria ephemerella (Lepidoptera, Pyralidae). Herbivory by Procambarus clarkti (Girard, 1852) (red swamp crayfish) was depressed in Habenaria repens Nutt. (aquatic orchid) by an endogenous ester (Wilson et al., 1999) and in Saururus cernuus L. (lizards-tail) by an array of lignoid metabolites (Kubanek et al., 2001). Rorippa nasturtium- aquaticum (L.) Hayek (syn. Nasturtium officinale) (watercress) 1s protected by 2-phenylethyl isothiocya- nate, a chemical synthesized by the endogenous glucosinolate-myrosinase system and highly toxic to freshwater gastropods in the genus Physella (Kerfoot et al., 1998: Newman et al., 1992). It is unknown whether E. densa used in our experiment utilizes chemical defenses such as these to prevent or reduce herbivory; however, the leaf structure and morphology of E. densa (rejected by P. insularum) are similar to that of the closely related and most-preferred H. verticillata, so it is unlikely that structural defenses were responsible for P. insularum’s rejection of E. densa. It is possible that E. densa possesses a chemically based feeding deterrent system to deter herbivory by P. insularum. While this question is beyond the scope of our experiment, it certainly merits further investigation. The majority of studies investigating the impact of herbivory by species of Pomacea have focused on P. canaliculata (channeled apple snail), the type species for the “canaliculata complex” of which P. insularum is a member. Cazzaniga (2002) suggested that any canali- culata-like apple snail had the potential to become a pest and cause damage to aquatic ecosystems, so it is likely that the field behavior of P. insularum will be similar to that reported for P. canaliculata. Snails belonging to the canaliculata complex have been introduced to other parts of the world as biocontrol agents to manage aquatic weeds (Cowie, 2002; Okuma et al., 1994; Wada, 1997). For example, Perera and Walls (1996) found that P. canaliculata effectively Page 253 controlled Pistia stratiotes L. (water lettuce) in the Caribbean. Unfortunately, the snails also feed on native plants, resulting in detrimental effects to the native fauna that rely on endemic plants for food and shelter (Simberloff and Stiling, 1996). Field experiments by Carlsson et al. (2004) revealed that P. canaliculata consumed most aquatic vegetation and caused bodies of water to become turbid with a dominance of planktonic algae. These workers also found that densities of >2 snails per m* in some of Thailand’s natural wetlands resulted in a shift in ecosystem state and function — virtually all aquatic plants were eaten and serious increases were recorded in nutrient concentrations and phytoplankton biomass (Carlsson et al., 2004). Herbiv- ory by P. canaliculata extends to commercially culti- vated crops as well; in fact, a density of 8 snails per m? can reduce yields by 90% in Oryza sativa L. (rice) (Naylor, 1996). Population densities of P. insularum- infested wetlands and flooded agricultural sites are often unavailable, but even small populations can explode quickly since canaliculata-type snails are highly fecund (Martin and Estebenet, 2002; Tanaka et al., 1999; Teo, 2004). Based on these reports and the results of our experiments, it is likely that wetland areas infested with P. insularum will experience severe damage similar to that reported by Carlsson et al. (2004). Also, drastically reduced yields should be expected in inundated agricultural crops (e.g., O. sativa, Colocasia esculenta (L.) Schott (taro)) grown in areas like Texas, Louisiana, Florida and Hawaii. These experiments provide additional documenta- tion regarding the feeding habits of P. insularum and the positive influence of moderate temperatures on herbivory and growth of the species. It is unlikely that macrophyte origin plays a role in P. insularum’s food selection, so special precautions must be followed to exclude this snail from ecosystems where aquatic macrophytes could be decimated by its presence. Many aquatic fauna rely on submerged vegetation as a habitat for nesting and spawning, so P. insularum’s indiscriminate consumption of aquatic macrophytes would have devastating consequences to Florida’s ecosystem. Other countries have attempted to use canaliculata-type snails to control nuisance aquatic plants, but their indiscriminate feeding habits have eliminated virtually all aquatic vegetation. Most aquatic systems support a variety of herbivores, but these species rarely feed at a level that significantly impacts macrophyte populations. Therefore, it is critically important that P. insularum and_ other canaliculata-type snails be excluded or eradicated in Florida and in other states at risk to prevent the decimation of critical aquatic ecosystems. Additional research should be conducted to assess the impact of P. insularum on other aquatic macrophytes, including floating, emergent and other submerged species. Of Page 254 particular interest would be predictive studies to determine the impact of population density of P. insularum on aquatic ecosystems. Acknowledgments. This research was supported by the Florida Department of Environmental Protection Bureau of Invasive Plant Management and by the Florida Agricultural Experi- ment Station. Mention of a trademark or a proprietary product does not constitute a guarantee or warranty of the product by the Florida Agricultural Experiment Station and does not imply its approval to the exclusion of other products that may be suitable. We thank Margaret Glenn for her contributions to this experiment. LITERATURE CITED BENSON, A. J. 2008. Pomacea insularum. USGS Nonindige- nous Aquatic Species Database, Gainesville, FL. Revision Date: 8/14/2007. http://nas.er.usgs.gov/queries/FactSheet. asp?speciesID=2599 CARLSSON, N. O. L., C. BRONMARK & L. A. HANSSON. 2004. Invading herbivory: The golden apple snail alters ecosystem functioning in Asian wetlands. Ecology 85(6): 1575-1580. CARLSSON, N. O. L. & J. O. LACOURSIERE. 2005. 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Tallahassee, FL. KERFOOT, W. C., R. M. NEWMAN & Z. HANSCOM. 1998. Snail reaction to watercress leaf tissues: reinterpretation of a mutualistic ‘alarm’ hypothesis. Freshwater Biology 40(2):201-213. KUBANEK, J., M. E. HAy, P. J. BROWN, N. LINDQUIST & W. FENICAL. 2001. Lignoid chemical defenses in the freshwa- ter macrophyte Saururus cernuus. Chemoecology 11:1-8. LACH, L., D. K. BRITTON, R. J. RUNDELL & R. H. COwIE. 2000. Food preference and reproductive plasticity in an invasive freshwater snail. Biological Invasions 2:279-288. MARTIN, P. R. & A. L. ESTEBENET. 2002. Interpopulation variation in life-history traits of Pomacea canaliculata (Gastropoda: Ampullariidae) in southwestern Buenos Aires Province, Argentina. Malacologia 44:153-163. NAYLOR, R. 1996. Invasions in agriculture: assessing the cost of the golden apple snail in Asia. Ambio 25:443-448. NEWMAN, R. M., Z. HANSCOM & W. C. KERFOOT. 1992. 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TANAKA, K., T. WATANABE, H. HIGUCHI, K. MryAMOTO, Y. YusaA, T. KIYONAGA, H. KryoTa, Y. SUZUKI & T. WADA. 1999. Density-dependent growth and reproduc- tion of the apple snail, Pomacea canaliculata: a density manipulation experiment in a paddy field. Research on Population Ecology 41:253—262. TEO, S. S. 2004. Biology of the golden apple snail, Pomacea canaliculata (Lamarck, 1822), with emphasis on responses to certain environmental conditions in Sabah, Malaysia. Molluscan Research 24(3):139-148. THOMPSON, F. G. 1997. Pomacea canaliculata (Lamarck, 1822) (Gastropoda, Prosobranchia, Pilidae): a freshwater snail introduced to Florida, U.S.A. Malacological Review 30:91. WADA, T. 1997. Introduction of the apple snail Pomacea canaliculata and its impact on rice agriculture. In: Proceedings, International Workshop on Biological Inva- sions of Ecosystems by Pests and Beneficial Organisms. National Institute of Agro-Environmental Sciences, Ministry of Agriculture, Forestry and Fisheries, Tsukuba, Japan. WILSON, D. M., W. FENICAL, M. Hay, N. LINDQUIST & R. BOLSER. 1999. Habenariol, a freshwater feeding deterrent from the aquatic orchid Habenaria repens (Orchidaceae). Phytochemistry 50:1333—1336. Instructions to Authors The Veliger publishes original papers on any aspect of malacology. All authors bear full responsibility for the accuracy and originality of their papers. Presentation Papers should include an Abstract (approximately 5% of the length of the manuscript), Introduction, Materials and Methods, Results, and Discussion. Short Notes should include a one- sentence Abstract. In taxonomic papers, all names of taxa must be accompanied by author and date of publication, and by a full citation in the bibliography. In papers on other subjects and in the non-taxonomic portions of taxonomic papers, author and date of names need not be accompanied by a full citation. 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LYONS’AND: MARTINUAVERY SNYDER. .205)05 eee 6 ocean tale ella ceca on eae eee 225 Injuries on Nautilus Jaws: Implications for the Function of Ammonite Aptychi ISABELLE KRUTA AND New Hi VANDMAN 3. 5: ciseee Gs sain neers er eee eee 241 Effect of Temperature and Feeding Preference on Submerged Plants by the Island Apple Snail, Pomacea insularum (@ Orbigny, 1839) (Ampullariidae) L. A. Gettys, W. T. Haier, C. R. MupGE AND T. J. KOSCHNICK ...........00000005 248 SMITHSONIAN INSTITUTION LIBRA MANOA A 3 9088 01444 | it | 1 IL (0 ay THE “VELIGER A Quarterly published by \ CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California R. Stohler (1901—2000), Founding Editor Volume 50 ISSN 0042-3211 December 16, 2008 CONTENTS A New Genus for Vesicomya inflata Kanie and Nishida, a Lucinid Shell Convergent with that of Vesicomyids, from Cretaceous Strata of Hokkaido, Japan KazuTakKa AMANO, RoBERT G. JENKINS, YUKITO KURIHARA AND STEFFEN KIEL ......... 255 A Quantitative Assessment of Spermatozoan Morphology in Nutricola confusa and Nutricola tantilla (Bivalvia: Veneridae). JAMES GERAGHTY, Micuaet P. RussELL AND NORMAN DOLLAHON .........-.00+0 0005 263 Effects of Estivation on the Concentrations of Selected Carboxylic Acids of Two Strains of Helisoma trivolvis Mark A. Brown, MicHaet J. CHEJLAVA, BERNARD FRIED, AND JOSEPH SHERMA ........ 269 Cenozoic Nacella (Patellogastropoda: Nacellidae) from Peru and Chile: Filling in the Gaps SIVELO MASH NID EN DRIES eueie cat tenet cies TOMA CDT Nace nna lacs WIENS an Oleh a aun a le ae 274 A New Species of Fissurella from Sao Pedro e Sao Paulo Archipelago, Brazil (Vetigastropoda, Fissurellidae) IGOR ZART CARD Ofer SIMONEC tis ce Pie eens ets cyetare ieee asec yaks teat elec dusce.ovel ei 292 The Gastropod Spiricella (Opisthobranchia: Umbraculidae) in the Recent Caribbean: A truly unexpected finding! CarLos MaRQUES DA SILVA & BERNARD M. LANDAU... 1.0... eee ees 305 Functional Anatomy of Bankia fimbriatula Moll & Roch, 1931 (Bivalvia: Teredinidae MARTA ULTAU IARTINS-SILVAVAND) WALTER) NARCHI) ac «i: 2 acces ca sc sem ee see eee 309 CoNnTENTS — Continued The Veliger (ISSN 0042-3211) is published quarterly by the California Malacozoological So- ciety, 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 His- tory, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. Number 4 THE VELIGER Scope of the Journal The Veliger is an international, peer-reviewed scientific quarterly published by the California Malaco- zoological Society, a non-profit educational organization. The Véeliger is open to original papers pertain- ing to any problem connected with mollusks. 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The Veliger 50(4):255—262 (December 16, 2008) THE VELIGER © CMS, Inc., 2007 A New Genus for Vesicomya inflata Kanie & Nishida, a Lucinid Shell Convergent with that of Vesicomyids, from Cretaceous Strata of Hokkaido, Japan KAZUTAKA AMANO Department of Geoscience, Joetsu University of Education, Joetsu 943-8512, Japan (e-mail: amano@juen.ac.jp) ROBERT G. JENKINS Faculty of Education and Human Sciences, Yokohama National University, Yokohama 240-8501, Japan YUKITO KURIHARA Department of Geology and Paleontology, National Museum of Nature and Science, Tokyo 169-0073, Japan STEFFEN KIEL Institute of Geosciences — Paleontology, Christian-Albrechts-University, 24118 Kiel, Germany Abstract. Newly collected specimens of the large bivalve Vesicomya inflata Kanie & Nishida from the lower Cenomanian Tenkaritoge Formation reveal that it is not a vesicomyid but is instead an unusual lucinid. The new monotypic genus Ezolucina is herein proposed for this species, which is characterized by venerid or vesicomyid shell shape, large size, a smooth surface, a deeply impressed lunule, one cardinal and one anterior lateral tooth in the right valve, and two cardinal teeth in the left valve. Stable carbon isotope analyses and petrographic observations show that the carbonate concretions yielding this species do not represent ancient hydrocarbon-seep deposits as was suggested previously. Rather, Ezolucina inflata and the associated solemyid, lucinid, thyasirid, and manzanellid bivalves lived in organic- and sulfide-rich sediment. INTRODUCTION Large fossil bivalves from the ““Middle Yezo Group” in northern Hokkaido, Japan, were described by Kanie & Nishida (2000) as Vesicomya inflata, and listed as being the earliest record for the genus Vesicomya (Campbell, 2006; Kiel & Little, 2006). They were found in two large calcareous concretions surrounded by mudstone, and were associated with the solemyid Acharax cretacea Kanie & Nishida, 2000 and the lucinid Miltha sp. Extant members of these bivalve taxa harbor chemoautotrophic endosymbionts, and because car- bonate concretions bearing these taxa have repeatedly been demonstrated to represent ancient hydrocarbon- seep deposits (Mayjima et al., 2005; Campbell, 2006), the concretions bearing Vesicomya inflata were interpreted as ancient hydrocarbon-seep deposits (Kanie & Nishida, 2000; Kanie et al., 2000). Living Vesicomya species have small shells that rarely exceed 13 mm in length (Cosel & Salas, 2001) and belong to a clade informally known as “small” vesicomyids, composed of the genera Vesicomya, Waisiuconcha, Isorropodon, Callogonia and Pliocardia (cf. Cosel & Salas, 2001; Krylova & Sahling, 2006). In contrast, the Cretaceous ““Vesicomya’”’ inflata reaches a length of up to 157 mm. In a recent revision of fossil North Pacific vesicomyids, Amano & Kiel (2007) pointed out that V. inflata has a deeply impressed asymmetrical lunule, a feature unknown in vesico- myids, and that its hinge structure had neither been described nor illustrated. Consequently, Amano & Kiel (2007) excluded V. inflata from the Vesicomyidae and suggested lucinid affinities instead. Newly collected specimens from the type locality of “Vesicomya’ inflata at Sanjussen-zawa Creek in northern Hokkaido possess hinge dentition, a pallial line, and adductor scars that clearly place this species in anew genus of the Lucinidae. In addition, petrographic thin sections and stable carbon isotope analyses of the fossil-bearing concretions are used to evaluate the Page 256 Nakagawa area Horokanai area \y Okhotsk Sea Vy \ Tappu area Lu ro) N. f & Yezo fore-arc mL = basin deposits Sample locality The Veliger, Vol. 50, No. 4 Figure 1. Type locality of Ezolucina inflata (Kanie & Nishida). environmental reconstruction of this site as an ancient hydrocarbon seep. MATERIALS AND METHODS The type material was examined at the Yokosuka City Museum, and nine new specimens of “‘Vesicomya”’ inflata were collected from four float carbonate concretions with molluscan fossils (HRK A—D) at the type locality in northern Hokkaido (Figure 1). Strati- graphically the specimens are from the My 4 Member of the Tenkaritoge Formation, which is considered early Cenomanian (Hashimoto et al., 1965; Nishida et al., 1998). The figured specimens and additional new material are housed at the Joetsu University of Education (JUE). The mineralogy of the fossil-bearing concretions was identified by thin-section observations and X-ray diffraction (XRD) analysis. Standard thin-section observations were performed by plane- and _ cross- polarized and reflected light microscopy. XRD analy- ses were carried out on unoriented slurries using a PANalytical X’Pert PRO at the Department of Earth and Planetary Science, the University of Tokyo (EPUT). Carbon and oxygen isotopes were analyzed using 2 to 10 mg powdered carbonate matrix. Carbon dioxide was produced from each powdered sample by reaction with 100% phosphoric acid in vacuo (25°C), and analyzed with a Finnigan MAT252 mass spec- trometer at EPUT. Carbon isotopic composition is expressed in the conventional 6 notation relative to the Vienna Peedee Belemnite standard (5'°C, %o vs. V- PDB, standard deviation <0.03%o). MINERALOGY AND ISOTOPE COMPOSI- TION OF CARBONATES Thin-section observation and XRD analysis show that the concretions are almost entirely composed of homogeneous micritic calcite and siliciclastic sediment. Structures typical for methane-induced carbonates, like clotted fabrics and stromatolitic laminae (cf. Greinert et al., 2002: Peckmann & Thiel, 2004), were not seen. d6'°C values range from —7.7 to —6.3%o (vs. V-PDB) Table 1 Stable carbon and oxygen isotope composition of the carbonate concretions from the type locality of Ezolucina inflata (Kanie & Nishida). Sample no. Texture Mineralogy 6bC 6°O HRKA micrite calcite —7.4 —1.3 HRK A micrite calcite —7.3 =I HRK B micrite calcite —6.3 —3.8 HRK C micrite calcite —6.7 —4.9 HRK C micrite calcite —6.5 AD) HRK C micrite calcite =e 5) K. Amano et al., 2007 (Table 1) and are also not indicative of anaerobic methane oxidation. Methane-derived carbonate usually shows 632C values lower than —40%o (cf. Peckmann & Thiel, 2004). Thus, neither thin section observations nor stable isotope analyses support the idea of Kanie & Nishida (2000) and Kanie et al. (2000) that “Vesico- mya’ inflata and associated mollusks lived at an ancient hydrocarbon seep. SYSTEMATICS Family Lucinidae Fleming, 1828 Genus Ezolucina Amano, Jenkins, Kurihara & Kiel, gen. nov. Type species: Vesicomya inflata Kanie & Nishida, 2000. Diagnosis: Large, inflated veneriform shell with smooth surface except for rough, low commarginal lamellae; posterior radial sulcus weak, and lunule deeply impressed. Hinge of right valve with one stout cardinal tooth and an anterior lateral tooth, left valve hinge with two cardinal teeth. Anterior adductor scar quadrate and anteriorly detached from pallial line; pallial line entire and deeply impressed. Discussion: Here Gabb, 1866 is similar to Ezolucina gen. nov. by having one cardinal tooth, one anterior lateral tooth in right valve and a deeply impressed lunule, but differs from Ezolucina by its smaller shell with fine ventral crenulations, and its much more deeply impressed lunule. Another large Cretaceous lucinid is Nipponothracia, which differs from Ezolucina by having an edentulous hinge and a very elongate anterior adductor scar (cf. Kanie & Sakai, 1997; Kase et al., 2007: Kiel et al., 2008). The medium-sized North American lucinid Nymphalucina Speden, 1970 from the Late Cretaceous Pierre Shale and Fox Hill Formation is oval in shape and lacks the sloping posterodorsal margin of Ezolucina. Trinitasia Maury, 1928, which was questionably placed in Lucinidae by Chavan (1969), is comparable in shell form and sculpture, but based on internal shell features Woodring (1982) showed that Trinitasia is not a lucinid but a mactrid. Etymology: A combination of the old name of Hokkaido (Ezo) and the genus Lucina. Ezolucina inflata (Kanie & Nishida, 2000) (Figures 2—9) Vesicomya inflata Kanie & Nishida, 2000: p. 79-82, JH, I, 2 Holotype: Articulated specimen, length 131.8 mm, height 105.4 mm, width76.3 mm, YCM-GP1173. Rage! sy Paratype: Articulated specimen, length 82.6 mm+, height 59.8 mm, width 35.8 mm+, YCM-GP1174. Topotypes: Articulated specimen, length 157.5 mm, height 123.8 mm, width 74.0 mm+, YCM-GP1177; right valve, length 33.7 mm, height 26.2 mm, JUE no. 15853; right valve, length 28.9 mm, height 21.9 mm, JUE no. 15854; articulated specimen, length 15.4mm, height 13.5 mm, width 5.9mm, JUE no. 15855. Type locality: Bed of Sanjussen-zawa Creek, 6.5 km upstream from where it flows into the Uryu River, Horokanai Town, Hokkaido (44°14'24’N, 142°5'26”E): My 4 Member of the Tenkaritoge Formation (locality R7203 by Nishida et al., 1998). Stratigraphic and geographic distribution: Late Creta- ceous (early Cenomanian); known only from the type locality. Original description: “Large shell (length [L] L = 130 mm in holotype and probably gerontic stage) probably of rounded triangular form (height [H] H/L = 0.8). Umbo (shell apex) situated almost centrally, 45-46% from anterior margin. Strongly inflated valve ([H)/breadth [B], H/B40.68). Shells of right and left are equivalved. Posterior end is truncated. In the juvenile to middle growth stage represented by paratype (L = 88.6 mm), the shell height is shortened (H/L = 0.68). Postero-dorsal end is truncated meeting with the posterior margin. Lunule and hinge form is the same as gerontic one. There is a lunule at antero-dorsal part. The ventral margin is weakly arched. Ligament probably external and long on the basis of morphology of the hinge area at postero-dorsal part. Test is very thick about 7 mm at the ventral margin. Shell surface 1s ornamented by concentric growth lines. The umbo bends strongly inside, where characteristic subumbonal pit exists at the inside of the shell apex.” Supplementary description: Shell large in size (up to 157.5 mm in length), thick, well inflated in adult specimens, veneriform, rather equivalve, slightly in- equilateral. Small specimens less inflated and Fela- niella-like in shape. Umbo projecting above dorsal margin, prosogyrate, situated anteriorly at two-fifths of shell length. Anterodorsal margin broadly arcuate, grading into rounded anterior margin; ventral margin broadly arcuate; posterodorsal margin straight, grad- ually sloping, forming blunt angle with subtruncated posterior margin. Surface smooth except for rough and low commarginal lamellae; coarse concentric ribs visible in right valve of a small specimen. Shallow radial sulcus running from beak to posterior end in large specimens. Hinge plate wide; right valve with one stout and triangular central tooth (3b), one elongate anterior Page 258 The Veliger, Vol. 50, No. 4 Figures 2-4. Type material of Ezolucina inflata (Kanie & Nishida). Figure 2. Holotype, length = 131.8 mm, YCM-GP1173. Figure 2a. Lateral view of right valve. Figure 2b. Dorsal view showing a posterior radial sulcus and deeply impressed lunule. Figure 3. Topotype, length = 157.5 mm+, YCM-GP1177. Figure 3a. Dorsal view. Figure 3b. Lateral view of left valve. Figure 4. Paratype, length = 82.6 mm, YCM-GP1174. Figure 4a. Lateral view of right valve. Figure 4b. Dorsal view showing a posterior radial sulcus and deeply impressed lunule. Figure 4c. Lateral view of left valve. K. Amano et al., 2007 Page 259 Figures 5-8. Additional specimens of Ezolucina inflata (Kanie & Nishida). Figure 5. Topotype, length = 28.9 mm, JUE no. 15854. Figure 5a. Hinge of right valve showing one strong cardinal and distinct anterior lateral tooth, length of hinge plate = 21.3 mm. Figure 5b. Dorsal view showing deeply depressed lunule. Figure 5c. Lateral view of right valve. Figure 6. Topotype, length = 39.9 mm+, JUE no. 15856. Figure 6a. Dorsal view of left valve showing deeply impressed lunule. 6b. Hinge of left valve showing two strong cardinal teeth, length of hinge illustrated here = 21.8 mm. 6c. Lateral view of left valve. Figure 7. Topotype, length = 15.4 mm, JUE no. 15855. Figure 7a. Lateral view of left valve. Figure 7b. Lateral view of right valve having rude concentric ridges. Figure 8. Topotype, length = 33.7 mm, JUE no. 15853. Figure 8a. Lateral view of right valve stressing anterior adductor scar. Figure 8b. Normal lateral view of right valve. lateral tooth (AI) parallel to hinge base, and a weak blunt node just below deeply impressed lunule; left valve with strong anterior cardinal (2), touching deeply impressed lunule, posterior tooth (4b) rather thin. Ligament occupying two-thirds of posterodorsal mar- gin. Anterior adductor scar elongate quadrate, moder- ate in size (adductor length = 9.1mm in JUE no. 15853; adductor length/shell length = 0.27) and about 75% of its length detached from pallial line (maximum distance from pallial line = 1.9 mm in JUE no. 15853); posterior adductor scar pear-shaped. Inner surface covered by coarse and distinct radial striations (Figure 9). Lunule broadly lanceolate, well defined, deeply impressed, slightly asymmetrical, slightly larger in right valve than in left valve, and occupying one- third of anterodorsal margin. Pallial line narrow, well- developed, entire, and quite distant from shell margin. Inner ventral margin smooth. Remarks: Very similar to Ezolucina inflata in shell shape and size is “Lucina’ colusaensis Stanton, 1895, a species that is apparently restricted to Upper Jurassic (Tithonian) to Lower Cretaceous (Hauterivian) cold- seep carbonates in northern California, USA (Stanton, 1895, p. 60, pl. 11, figs. 4, 5; Campbell, 2006; Kiel et al., 2008). Compared to our material of Ezolucina inflata the thickness of ‘“Lucina’ colusaensis resembles that of the less inflated specimens of Ezolucina inflata. Unfortunately, ‘Lucinda’ colusaensis is usually poorly preserved and features of the interior of this shell are unknown (Stanton, 1895; SK, pers. observation; K.A. Campbell, personal communication 2007); thus a Page 260 The Veliger, Vol. 50, No. 4 Figure 9. Muscle scars and pallial line of Ezolucina inflata (Kanie & Nishida) based on JUE no. 15853. Abbreviations: aa, anterior adductor scar; pa, posterior adductor scar; pl, pallial line. confirmation of its generic position must await the availability of well-preserved specimens. Another large lucinid with prominent umbos is Saxolucina (Megaxinus) matsushitai Matsumoto (1971, p. 663-665, pl. 1, fig. 1, pl. 2, figs. 1-3) from the Oligocene Setogawa Group [now considered as Miocene in age; see Watanabe (1988)] in central Japan, but this species is clearly distinct from Ezolucina inflata by having a less inflated shell with an edentulous dentition. Ezolucina inflata resembles Here excavata (Carpen- ter, 1857) in having a deeply depressed lunule, one cardinal tooth and anterior lateral tooth in the right valve, and two cardinal teeth in the left valve. However, Here excavata differs from Ezolucina inflata by its subcircular shell with concentric lamellae, many fine ventral crenulations and more deeply sunken lunule. Its veneriform shape with the strongly sloping postero- dorsal margin sets Ezolucina inflata apart from most other Cretaceous lucinids, which mostly have a nearly round outline (e.g., Discoloripes septentrionalis Kelly, 1992; “Lucina’ spp. in Stephenson, 1952; Callucina olea Vokes, 1946; “‘Lucina’ aquensis Holzapfel, 1889, p. 188, pl. 19, fig. 4). A lucinid with similar hinge dentition was described and figured as Lucina sub- nummismalis dOrbigny, 1850 from the Campanian Vaals Greensand of Germany (Holzapfel, 1889, p. 187— 8, pl. 19, figs. i—3). This species is distinct from Ezolucina inflata because it is very flat, has distinct commarginal ribs, and lacks the strongly sloping posterodorsal margin of Ezolucina inflata. Also ‘‘Luci- na aff. valentula de Lor.” described by Ascher (1906, p. 161, pl. 14: 5) from a potential Hauterivian seep site in eastern Czech Republic lacks the strongly sloping posterodorsal margin of Ezolucina inflata. DISCUSSION Examination of the type and additional specimens clearly shows that Vesicomya inflata is a member of Lucinidae because it has the hinge structure and the adductor muscle scar and pallial line patterns typical of Lucinidae (1.e., a lucinoid hinge dentition, a broadly lanceolate, asymmetric, sunken lunule, and an elongate anterior adductor muscle scar detached from the pallial line). The previous assignment of this species to the Vesicomyidae was due to the superficial resemblance in shell outline and the lack of the information on the shell interior. The new monotypic genus Ezolucina based on V. inflata is more elongate than most lucinids and has moderately prominent umbones somewhat suggestive of a vesicomyid or an eomiodontid. How- ever, shells of these families can easily be distinguished from those of the Lucinidae by their hinge and muscle scar patterns. In their compilation of the stratigraphic ranges of mollusks at cold seeps, Kiel & Little (2006) listed V. inflata as the oldest fossil occurrence of Vesicomya, based on the available literature (Kanie & Nishida, 2000). When Amano & Kiel (2007) subsequently revised the North Pacific fossil record of the Vesico- myidae, they could not confirm any of the previous records of this genus. One of the oldest “‘small” vesicomyids related to Vesicomya (cf. Cosel & Salas, K. Amano et al., 2007 Page 261 Table 2 Faunal list based on fossils that we collected from the type locality of Ezolucina inflata (Kanie & Nishida). All of these species possibly harbor symbiotic bacteria. Numbers indicate the number of recovered individuals, capital letters are sample designations. Species Carbonate no. (HRK -) A By (Ce 1B) Acharax cretacea Kanie and Nishida D) Nucinella ? sp. 1 Ezolucina inflata (Kanie and Nishida) 3 2 4 Miltha? sp. 1 Thyasira sp. 1 2001; Krylova & Sahling, 2006) is “Vesicomya”’ kawadai (Aoki, 1954) from the lower Miocene Taira Formation in Fukushima Prefecture, Japan (Aoki, 1954; Kamada, 1962), but more material is required to confirm its generic position. Another certain record of a “small” vesicomyid is /sorropodon frankfortensis Amano & Kiel, 2007 from the lower Miocene Astoria Formation in Washington State, USA. Kanie et al. (2000) suggested that the carbonate concretions from Sanjussen-zawa Creek were ancient cold-seep deposits because of the presence of chemo- symbiotic species like solemyids, vesicomyids, and lucinids. Even when the vesicomyids are removed from this list, most taxa from these concretions (see Table 2) rely at least partly on symbiotic sulfur-oxidizing bacteria. However, the isotope data and the lithological observations presented here do not support a recon- struction as ancient cold-seep deposit. The data only suggests that these taxa lived in organic- and sulfide- rich sediment favorable for species with sulphophilic symbionts. Three of the five taxa from the concretions at Sanjyussen-zawa Creek (Acharax, Thyasira, and Miltha) can also be found in the mudstone and siltstone of the Yezo Group (Tashiro, 2004) and their relative abundance at this site might be due their early diagenetic preservation in the concretions. The exceptionally large size of Ezolucina inflata is remarkable; among Cretaceous lucinids it even exceeds that of the cold-seep restricted Nipponothracia. Among Recent species, it matches the size of Meganodontia acetabulum Bouchet & Cosel, 2004, recently described as the largest living lucinid from a depth of 256-472 m. The species was found in an area of presumed diffuse gas seepage and was associated with other bivalves bearing chemotrophic endosymbionts, like solemyids, thyasirids, and other lucinids. This set of taxa is similar to that associated with Ezolucina inflata, but our carbon isotope data do not indicate gas seepage in its environment. Most other modern deep-water lucinids rarely exceed 50 mm in length (Cosel, 2006) and are thus much smaller than Ezolucina inflata. However, deep-water environments of the Late Cretaceous were inhabited by a number of exceptionally large taxa like bivalve Inoceramus or the capulid limpet Gigantocapu- lus giganteus (cf. Takahashi et al., 2007). Thus, the paleoenvironment of Ezolucina inflata remains enig- matic. Acknowledgments. We are very grateful to James L. Goedert (Burke Museum, Seattle) for his critical reading of this manuscript. We thanks John D. Taylor (Natural History Museum) and Geerat J. Vermeij (University of California at Davis) for their review. We also thanks Tamio Nishida (Saga University) for his offering some fossil specimens. LITERATURE CITED AMANO, K. & S. KIEL. 2007. Fossil vesicomyid bivalves from the North Pacific region. Veliger 49:270-293. AOKI, S. 1954. Mollusca from the Miocene Kabeya Forma- tion, Joban coal-field, Fukushima Prefecture, Japan. Science Reports of the Tokyo Kyoiku Daigaku, Section C 3:23-41. ASCHER, E. 1906. Die Gastropoden, Bivalven und Brachio- poden der Grodischter Schichten. Beitrage zur Palaonto- logie und Geologie Osterreich-Ungarns und des Orients 19:135-172. BOUCHET, P. & R. VON COSEL. 2004. The world’s largest lucinid is an undescribed species from Taiwan (Mollusca: Bivalvia). Zoological Studies 43:704-711. BRETSKY, S. S. 1976. Evolution and classification of the Lucinidae (Mollusca; Bivalvia). Palaeontographica Amer- icana 8:219—337. CAMPBELL, K. A. 2006. 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U.S. Geological Survey Professional Paper 306-F:541—759, pls. 83-124. The Veliger 50(4):263—268 (December 16, 2008) THE VELIGER © CMS, Inc., 2007 A Quantitative Assessment of Spermatozoan Morphology in Nutricola confusa and Nutricola tantilla (Bivalvia: Veneridae) JAMES GERAGHTY, MICHAEL P. RUSSELL* AND NORMAN DOLLAHON Department of Biology, Villanova University, Villanova, PA 19085, USA (e-mail: michael.russell@villanova.edu) Abstract. The brooding bivalves, Nutricola tantilla and N. confusa overlap in their geographic distributions, habitats, and modes and timing of reproduction. Based on previous studies we infer that males of both species release spermatozoa into the water column; while females retain developing embryos in a brood chamber. Females release fully formed juveniles and there is no pelagic larval stage. We hypothesized that the muco-ciliary processes of particle selection and retention may act on differences in sperm morphology and contribute to reproductive isolation. We extracted sperm cells from both species and quantified nine linear measurements: the lengths of the acrosome, nuclear, midpiece and tail regions, and five different width measurements. We found significant differences in the lengths of the acrosome, midpiece, and tail. We also found that N. confusa produces dimorphic sperm and this is the first report of sperm dimorphism in the Veneroidea. Despite the significant differences in lengths, it is likely that other prezygotic mechanisms are responsible for reproductive isolation. INTRODUCTION Nutricola tantilla and N. confusa are morphologically similar, small bivalves (<10 mm shell length), that inhabit the top 2.5 cm of soft substrata in the intertidal to shallow-subtidal zones of protected bays of western North America (Coan et al., 2000). The reported geographic range of Nutricola tantilla is from Prince William Sound, Alaska to Isla Cedros, Baja California and N. confusa occurs from Coos Bay, Oregon to Carmel Bay, California. Where their ranges overlap they are sympatric and both species are very common in Bodega Bay, California (Grosholz & Ruiz, 1995). In earlier studies, the two species have been referred to as Transennella and there is some disagreement over the taxonomy (Lindberg, 1990). Females of both species are generally larger than males (Hansen, 1953; Asson-Bartres, 1988; Russell and Huelsenbeck, 1989). Hansen (1953) performed histo- logical examinations of 371 specimens and found “5 were in a state of reversal from male to female” (p. 319). Some studies have cited this work as evidence for protandry (Kabat, 1985, 1986; Asson-Bartres, 1988) whereas Mottet (1988) attributes the size disparity to differential growth rates. Both species lack pelagic larval stages and females brood their developing embryos and early juvenile stages in a pouch located between the inner demibranch and visceral mass (see figures 2 and 3 in Kabat, 1985 for detailed SEMs). Broods can be found throughout the year but there is seasonal variation in reproduction * Correspondence and reprints request with higher levels during the summer and fall (Asson- Batres, 1988; Russell & Huelsenbeck, 1989). During peak periods of reproduction brood number can be as high as 300 and is a function of female size (Kabat, 1985; Russell & Huelsenbeck, 1989). Sperm storage has not been reported in Nutricola (Mottet, 1988), we have not observed it, and therefore conclude that these species outcross. We infer that males release sperm into the water column because individuals produce only eggs or sperm at any one time and females retain their eggs for brooding. Sperm probably enter the mantle cavity of a female through the siphons. This fertilization mechanism, called “spermcast mating” (Bishop & Pemberton, 2006), has been proposed for other outcrossing brooding bivalves (Oldfield, 1964; Sellmer, 1967) and shown to be the method of sperm transfer for the brooding bivalve, Mysella tumida (O’Foighil, 1985b). Once inside the mantle cavity, the mechanics of directing sperm dorsally to the ovaries and unfertilized eggs is unknown but could involve chemotaxis and/or selective muco- ciliary activity of the gills. Nutricola tantilla and N. confusa are sympatric, reproduce at the same time of year, and presumably females are exposed to sperm released from males of both species. One question that arises from these circumstances is how is reproductive isolation main- tained? The purpose of our study was to quantify the gross morphology of spermatozoa from N. tantilla and N. confusa to determine whether differences in size/ shape could contribute to the maintenance of repro- ductive isolation. This study represents the first attempt Page 264 The Veliger, Vol. 50, No. 4 Lengths Tail Widths 5 6 7 3 © Figure 1. Schematic diagram of spermatozoa found in Nutrico/a illustrating the nine linear dimensions measured. Lengths: 1 = acrosome, 2 = nuclear region, 3 = midpiece, 4 = tail. Widths: 5 = acrosome midpoint, 6 = acrosome juncture with nuclear region, 7 = nuclear region midpoint, 8 = nuclear region juncture with midpiece, 9 = midpiece. to quantify Nutricola spermatozoa morphology using electron microscopy. METHODS AND MATERIALS Samples of both species of clams were collected from Bodega Bay California on March 26, 2007, shipped overnight to Villanova University, separately main- tained in a sea table (10°C and 32%o), and fed a mixture of phytoplankton cultures of Tetraselmis sp., Thalas- siosira sp., Isochrysis galbana, and Chaetocerous muelleri until processed. Attempts to induce spawning with thermal shocking methods (Castagna & Kraeuter, 1981; Deming & Russell, 1999) were unsuccessful so we resorted to extracting sperm via gonad squashes. Individuals of each species were dissected in separate containers of sea water. The gonads were removed, gently macerated, and released sperm in the sea water. Samples of the seawater were examined with a compound microscope for the presence/activity of spermatozoa. When active spermatozoa were identified, additional seawater sam- ples with sperm were pipetted on to ploy-L-lysine coated cover slips. The sperm were fixed at 4°C in 2.5% glutaraldehyde in 0.1 M cacodylate buffer, adjusted to pH 7.4 and 900 mOsm with 0.4 M NaCl and 2 mM CaCl. After a rinse in the same buffer, samples were postfixed in 1% osmium tetroxide, in the same buffer, and rinsed again. Samples were dehydrated in an ascending series of ethanol dilutions (25%, 50%, 75%, 95%, and 100%) and critical point dried using CO; as transitional fluid. Cover slips were sputter coated with gold/palladium and observed in a Hitachi S-570 SEM at 5 kV. Digital images of intact spermatozoa were captured and then measured using ImageJ software version 1.37 V (Abramoff et al., 2004). Nine separate linear measurements were recorded on each intact spermato- zoan (Figure 1): lengths of the acrosome, nuclear region, midpiece and tail; and widths of each of the three head regions (acrosome, nuclear, and midpiece) as well as the widths of the boundaries between adjacent head regions. All data were tested for normality using a Shapiro- Wilk W test and the nine linear dimensions between species were compared using a t-test when both data sets were normally distributed, or a Wilcoxon Rank Sum test when either (or both) data sets were not normally distributed. A Principle Component Analysis (PCA) was used to combine all morphometric data to visualize the degree of separation between the two species based on spermatozoa morphology. Finally, a discriminant analysis was performed to assess how many of the samples would be correctly assigned to each species based on the nine linear measurements. All J. Geraghty et al., 2007 Page 265 Figure 2. Examples of spermatozoa found in Nutricola. The scale bars = 10 um. A. Nutricola tantilla. B. Nutricola confusa. C. This type of sperm representing a distinct morphology (rounded-head) was found only in N. confusa. analyses were performed using JMP (Version 4.04, SAS Institute Inc.). RESULTS Active sperm were found in all samples of male clams; 3 males were found for N. tantilla and 2 males for N. confusa. Electron microscopy preparations were pro- cessed for each male and intact spermatozoa were identified and measured for both N. tantilla (n = 13) and N. confusa (n = 18). Both species exhibited markedly elongated heads and examples of these cells illustrating the three distinct regions of the head are shown in Figure 2. Furthermore, a morphologically different sperm with a round head was found only in samples from N. confusa (Figure 2C). Significant differences in the lengths between the species were found in the acrosome (Z = 2.62, P = .0087), tail (Z = 3.70, P = .0002), and midpiece (t = 4.39, P = .0001) regions. In all three cases the spermatozoa of N. tantilla were significantly longer than N. confusa (Figure 3). No significant difference was found in any of the width measurements or the length of the nuclear region. The range-frame box and whiskers plots show that although the spermatozoa from WN. tantilla are longer, there is considerable overlap with N. confusa (Figure 3). The results of a PCA are displayed in Figure 4 and a Page 266 60 - 50 F = * of 4] Length (um) 3 T Acrosome Tail The Veliger, Vol. 50, No. 4 3.5 ab =| cH Nuclear Midpiece EJ N. confusa Width (um) S Paras Acrosome (midpoint) Acrosome/ Nuclear (junction) Nuclear (midpoint) | N. tantilla 4 55 Nuclear/ Midpiece (junction) Midpiece (midpoint) Figure 3. Box and whiskers plots of the nine linear measurements of N. tantilla (shaded) and N. confusa (open) spermatozoa. The horizontal lines within each box are the medians, the edges of the boxes are the 25" and 75" percentiles and the whiskers are the 5" and 95" percentiles. Significant differences (*) were found in the acrosome, tail, and midpiece lengths. discriminant analysis correctly identified 87% (27 out of 31) of the measured spermatozoa. DISCUSSION Both species showed spermatozoa with elongated heads composed of three distinct regions (Figure 2). The designations of acrosome, nuclear, and midpiece (Figure 1) are based on the relative positions of these regions in the sperm of other taxa (Franzén, 1956) and comparison with the illustration and description of the spermatozoa of Transennella (= Nutricola) tantilla in (Thompson, 1973). The mean lengths of the acrosome, nuclear, and midpiece regions of N. tantilla from our samples are 13.68, 2.47, and 1.27 (tm) respectively, which are comparable to the lengths Thompson (1973) reported: 15.0, 2.4, and 1.0 (um). Franzén (1983) noted that the acrosome 1s a “prominent structure” in bivalve spermatozoa (as is the case here) and commented on Thompson’s (1973) description of N. tantilla sperma- tozoa that “‘in spite of its unusual proportions [it] seems to belong to the primitive type.” Spermatozoa with a distinctly different morphology were found only in N. confusa (Figure 2C). We did not observe any intermediate stages between the “round headed”’ sperm and the mature sperm with the elongated acrosomes (Figure 2B). This observation strongly suggests the presence of sperm dimorphism in N. confusa. Sperm dimorphism is relatively uncom- mon in bivalves having been reported in only a few species (Ockelmann, 1965; Jespersen et al., 2001; Jespersen & Liitzen, 2001; Litzen et al., 2001; Jespersen et al., 2002; Jespersen et al., 2004; Litzen et al., 2004). We found this second type of sperm in all of the SEM preparations of N. confusa and in none of the preparations from WN. tantilla. This finding 1s the first reported case of sperm dimorphism in the Veneroidea. Other reports of sperm dimorphism occur in the Galeommatoidea where one species in one genus can exhibit sperm dimorphism, e.g., Kurtiella bidentata (as J. Geraghty et al., 2007 Page 267 3 PC1 Figure 4. Results of Principle Component (PC) analysis using the nine measures illustrated in Figure 1. Nutricola tantilla (shaded, n = 13) and N. confusa (open; n = 18). Mysella), and another closely related species does not, e.g., M. tumida (Ockelmann & Muus, 1978; O’Foighil, 1985a). It appears that this is the situation with the congeners N. tantilla and N. confusa. The acrosome, midpiece and tail of the Nutricola tantilla sperm are significantly longer than those of N. confusa but no differences were found in the length of the nuclear region or any of the five width measure- ments (Figure 3). There is a significant difference in the overall mean lengths of spermatozoa — N. tantilla 65.26 um + 4.64 and N. confusa 56.93 um + 5.73 (+ SD, Z = 3.78, P = .0002). Although there are significant morphological differences between the sperm in these species there is also considerable overlap in the variables measured. This point is illustrated by the PCA plot (Figure 4) which shows a limited degree of separation between the two species. There are at least three hypotheses for the functional significance of elongated sperm heads in bivalves. Recently, Jespersen & Litzen (2007) proposed that this morphology allows sperm cells to better circum- vent retention by the gills thus facilitating fertilization. Franzén (1983) found that elongated sperm heads are correlated with larger eggs and may aid in sperm penetration. Finally, Jespersen et al. (2001) proposed that the elongated sperm heads of the euspermatozoa of Pseudopythina macrophthalmensis may promote storage of sperm in seminal receptacles. Neither N. tantilla nor N. confusa have seminal receptacles and do not store sperm so the later hypothesis does not apply to these species. However, the unusually long sperm heads of N. tantilla and N. confusa could function in either gill circumvention or penetration of the large lecithotrophic eggs. The study of the muco-ciliary processes of particle selection and retention in bivalves has a long and rich history (see Ward & Shumway, 2004 for review). The focus of these studies has been on feeding biomechanics and the ecological role bivalves play in benthic-pelagic coupling processes. During preingestive processing, “there are opportunities for particle selection based upon quantitative and qualitative aspects of the particles” (Ward & Shumway, 2004:85). Spermatozoa cells of spermcast-mating, brooding bivalves like Nutricola, are within the size-range of particles selected via the muco-ciliary processes (Mohlenberg & Riisgard, 1978) and are likely subject to these processes. The differences in sperm morphology demonstrated here while significant, are probably insufficient by them- selves to account for species-specific spermatozoa recognition. These species produce hundreds of eggs compared to the hundreds of thousands produced by broadcast spawning taxa and cannot afford postzygotic isolation. Therefore it is likely that factors other than spermatozoa morphology play a role in maintaining reproductive isolation via prezygotic mechanisms. Page 268 Acknowledgements. We thank Ted Grosholz of UC Davis for kindly supplying samples from Bodega Bay and sharing his extensive knowledge of Nutricola biology. He also provided excellent advice on bivalve husbandry. We appreciate the careful and detailed constructive criticism provided by two anonymous reviewers. We are grateful for the laboratory assistance of S. Shrom, L. Bloch, C. Hynes, L. Elliot, C. Kitchell, and V. Garcia. The Biology Department at Villanova supported this work and the data formed the basis of J. Geraghty’s senior thesis. M. P. Russell was supported by the Ocean Sciences Division Biological Oceanography of NSF. LITERATURE CITED ABRAMOFF, M. D., P. J. 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HUELSENBECK. 1989. Seasonal variation in brood structure of Transennella confusa (Bivalvia: Veneridae). The Veliger 32:288—295. SELLMER, G. P. 1967. Functional morphology and ecological life history of the gem clam, Gemma gemma (Eulamelli- branchia: Veneridae). Malacologia 5:137—223. THOMPSON, T. E. 1973. Euthyneuran and other molluscan spermatozoa. Malacologia 14:167—206. WARD, E. J. & S. E. SHUMWAY. 2004. Separating the grain from the chaff: particle selection in suspension- and deposit-feeding bivalves. Journal of Experimental Marine Biology and Ecology 300:83—130. THE VELIGER 2) The Veliger 50(4):269-273 (December 16, 2008) © CMS, Inc., 2007 Effects of Estivation on the Concentrations of Selected Carboxylic Acids of Two Strains of Helisoma trivolvis MARK A. BROWN,' MICHAEL J. CHEJLAVA,! BERNARD FRIED** AND JOSEPH SHERMA' ‘Department of Chemistry and *Department of Biology, Lafayette College, Easton, Pennsylvania 18042, USA Abstract. High-performance ion exclusion column liquid chromatography was used to analyze the effects of estivation on certain carboxylic acids in the digestive gland-gonad complex (DGG) of a Pennsylvania (Pa) and Colorado (Co) strain of Helisoma trivolvis. The DGG samples were extracted using 50% Locke’s solution, followed by cleanup using anion-exchange solid phase extraction and analysis by ion exclusion high performance liquid chromatography with ultraviolet detection. Succinic, pyruvic, malic, and fumaric acids were detected and quantified in the DGGs of both estivated and unestivated snails at concentrations ranging from 0.91 to 2200 ppm. There was a significant (Student’s f-test, P = 0.05) reduction in the concentrations of succinic, pyruvic, and malic but not of fumaric acid in unestivated versus estivated Helisoma trivolvis (Pa). For Helisoma trivolvis (Co), there was a significant increase in succinic but not in fumaric, pyruvic, or malic acids in unestivated versus estivated snails. The reduction of certain carboxylic acids in the DGG of Helisoma trivolvis (Pa) suggests that estivation stimulates a decreased production of these acids or increased utilization by the snail tissue. Differences in the concentrations of certain acids between H. trivolvis (Pa) and H. trivolvis (Co) probably reflect strain differences. INTRODUCTION Estivation (also written as aestivation) of pulmonate snails in the family Planorbidae relates to a dormancy of these snails under conditions of drying out. Estivation allows the snails to survive long periods of drought. Morevoer, snails infected with larval trema- todes may retain their infections during estivation and transfer of the larval parasites to new hosts may occur when the snails emerge from estivation following submersion in fresh water. In the field, such estivation occurs when snails are subjected to drying conditions in lakes or ponds during short term or extended periods of drought. In the laboratory, estivation can be induced by subjecting the planorbids to a high relative humidity (circa 98%) and temperatures of about 22 to 24°C ina moist closed chamber as described by White et al. (2007). During estivation, snail metabolism is reduced and the organisms do not feed on exogenous food stuff. Information at the cellular and molecular level of the effects of estivation on planorbids is sparse. In our laboratory, we have begun estivation studies on planorbids in the genera Biomphalaria and Helisoma. Snails in these genera are important vectors of larval trematodes and nematodes and are associated with the transmission of numerous helminthic diseases to humans and wildlife; planorbid snails in these genera also serve as models for various research studies in the * Corresponding author, e-mail: friedb@lafayette.edu biomedical sciences. Our studies to date on the topic have examined the effects of experimentally-induced estivation on various analytes in both uninfected and infected planorbids in the genera Biomphalaria and Helisoma. In general, our snail estivation studies have demonstrated a reduction of most of the analytes we examined and in most cases parasitism by larval trematodes has exacerbated the effects of estivation on diminution of certain metabolites in the snails. The studies reported in this paper are a continuation of the effects of estivation on analytes in two strains of planorbid snails in the genus Helisoma. More informa- tion on the rational for the present work is given in the paragraphs below. We maintain two strains of Helisoma trivolvis (Say, 1816) in our laboratory, one of which is H. trivolvis (Co) and the other H. trivolvis (Pa). The Co strain lacks melanin, is refractory to infection with miracidia of all trematodes tested to date, and is easy to culture in the laboratory (Schneck & Fried, 2005). It is also used as a model in invertebrate neurobiology (Kater, 1974). The Pa strain is ubiquitous in lakes and ponds in North America, is pigmented with melanin, and is infected with various species of larval trematodes (Schmidt & Fried, 1997). Recent studies in our laboratory have examined the effects of estivation on various analytes in Biomphalaria glabrata (Say, 1816). Studies on lipids (White et al., 2006), carbohydrates (Jarusiewicz et al., 2006) and lipophilic pigments (Arthur et al., 2006) in the DGGs Page 270 The Veliger, Vol. 50, No. 4 of these snails showed a decrease in these analytes as a function of snail estivation. One study on carboxylic acids in B. glabrata has determined changes in certain acids as a function of infection with Schistosoma mansoni larvae (Massa et al., 2007). One study on B. glabrata showed alterations in the concentrations of certain carboxylic acids as a function of estivation (Bezerra et al., 1999). Detailed studies on effects of estivation on the carboxylic acid content of any strain of H. trivolvis are not available. Therefore, the purpose of this study was to determine the effects of estivation on certain carboxylic acids in both the Pa and Co strains of H. trivolvis. MATERIALS AND METHODS Snail Maintenance H. trivolvis (Co) has been maintained in our laboratory in continuous culture since the mid 1980s. This strain is maintained in Mason jar cultures, 15 to 20 snails per jar, in 800 mL of aerated artificial spring water (ASW). For further details, including the formulation of the ASW, see Schneck & Fried (2005). H. trivolvis (Pa) is available from April through November from local farm ponds and lakes in North- ampton County, Pa (see Schmidt & Fried, 1997 for details). H. trivolvis (Pa) can be collected in the wild, brought into the laboratory, and maintained there for several months using the same cultivation procedure described for H. trivolvis (Co). Because some of these snails may be naturally infected with larval trematodes, they were examined by routine snail isolation proce- dures for larval trematodes; infected snails were removed from cultures and discarded. Estivation Usually 5 to 15 H. trivolvis (Pa) snails of each strain were estivated for 2, 3, or 7 days and a similar number of H. trivolvis (Co) for 7 days in a moist chamber at 24°C and a relative humidity of 98%. Details of the estivation chamber design were given in White et al. (2006). Preliminary studies showed that H. trivolvis (PA) did not survive the effects of estivation as well as H. trivolvis (Co), and therefore we used shorter estivation times for the studies on H. trivolvis (Pa). At the end of each estivation period, snails were tested for survival by immersing them in ASW. Live snails became activated within 0.5 hr in ASW as shown by the extension of the head foot through the aperture. Snails that did not extend the head foot through the aperture were examined by mechanical probing with a needle after the shells were removed. Those that were not responsive were considered dead. The number of snails that survived estivation was recorded. Controls (unestivated snails) were maintained in Mason jar cultures and fed leaf lettuce as described above for the same times as those that estivated. Sample Preparation Each snail was removed from the ASW and placed in a Petri dish. The shell was cracked gently and removed from the snail body. The DGG was dissected from the body and homogenized with 6 mL of 50% Locke’s solution with a glass homogenizer. The homogenizer was washed with 2—3 mL of solution, which was then added to the homogenate. The DGG homogenate was centrifuged at 2500 g for 10min at 25°C. The carboxylic acids were recovered from the supernatant by solid-phase extraction (SPE) as described below. Each sample represented the supernatant of one DGG and had a final volume of 8 + 1 mL. Carboxylic Acid Extraction SPE was performed as outlined by Massa et al. (2007a). The acids were extracted from the DGG homogenate using Varian strong anion exchange columns (quaternary amine; 100 mg; 3 mL, Varian Inc., Palo Alto, Ca, USA). Under vacuum, the columns were cleaned and activated with 1 mL of 0.5 M HCL, 1 mL of methanol, and 2 mL of deionized (DI) water. The DGG homogenate supernatant was then passed through a column under vacuum. The column was cleaned again with 2 mL of DI water. The carboxylic acids were eluted from the columns using 1 mL of 0.5 M sulfuric acid. HPLC Analysis Acetic, fumaric, lactic, malic, pyruvic, and succinic acid salts were purchased from Sigma (St. Louis, Mo, USA). Stock solutions of each organic acid were prepared at a concentration of 1.00 x 10°? ppm in 0.5 M H>SO, and the stock solutions were diluted to 10, 25, 50, and 100 ppm. High performance liquid chromatography (HPLC) was performed at 30°C using an Agilent Technologies (Wilmington, DE, USA) 1100 Series HPLC Instrument with an autosampler amd ultraviolent (UV) detection at 210 nm. A Bio-Rad Laboratories (Hercules, CA, USA) Aminex ion exclusion HPX-87H column (300 X 7.8 mm) was used. 0.5 mM sulfuric acid was used as the mobile phase with an injection volume of 100 uL. Linear calibration curves were generated using Microsoft Excel relating standard concentrations to their peak areas. The interpolated amounts of each organic acid quantified by HPLC were calculated using the following equation: M. A. Brown et al., 2007 Page 271 Table 1 Carboxylic acid concentrations in the digestive gland-gonad complex (DGG) of unestivated and estivated H. trivolvis (Pa). Unestivated DGG Estivated DGG (2 Days) Acid Sample Size ppm (ug/g + Standard Error) Sample Size ppm (ug/g + Standard Error) Fumaric 18 100 + 8.0 11 9622 18:5 Malic 9 450 + 85 5) 290 + 53 Pyruvic 8 2X) S= Sei 4 Sya= PX) Succinic 10 350 + 66 6 540 + 180 Unestivated DGG Estivated DGG (3 Days) Acid Sample Size ppm (ug/g + Standard Error) Sample Size ppm (ug/g + Standard Error) Fumaric 16 130 + 18 fl 140 + 13 Malic 9 DIO 35S 7 <2.0* Pyruvic 10 23 + 7.4 af <2.0* Succinic 9 280 + 36 7 <2.0* Unestivated DGG Estivated DGG (7 Days) Acid Sample Size ppm (ug/g + Standard Error) Sample Size ppm (ug/g + Standard Error) Fumaric 8 120 + 8.8 4 98 + 16 Malic 7 350 + 54 4 = < posterior Figure 10. Position of apex in Spiricella unguiculus Rang & Des Moulins, 1828 vs. Spiricella redferni n. sp. European Cenozoic basins and in the Northeastern Atlantic and Mediterranean (Silva & Landau, 2007, with references). Still nothing is known of the ecology of this genus, and no living animal or soft parts have ever been found. Unfortunately, these new finds add no new information concerning the ecology of Spiricella, as the shells were found in samples of beach drift from the extreme southwest tip of Abaco, between the locality of Sandy Point and the promontory of Rocky Point. Rocks border the beach, with Thalassia meadows beyond the rocks. This shell grit contains an assemblage of shells from all the neighboring environments: rocky and sandy substrates, Thalassia meadows, as well as deeper water environments, which lie close by. As pointed out by Silva & Landau (2007), and almost every other researcher struggling to make sense of this genus, problems are posed by the paucity of specimens; fossil and Recent, lack of knowledge of the soft parts, and lack of distinctive shell characters. The eastern shells are fairly uniform in shape, all more or less rectangular, with length/width ratio of 22.22. Only the presumably immature Recent shell from Serini, Mauritania is less elongated, with a ratio of 1.79 (Geuze & Hoeksema, 1994). All the shells have a smooth paucispiral proto- conch of 1.25—1.75 whorls. The position of the apex is eccentric in all, placed about 1/5 distance from the posterior edge and to the left (Figure 10). The Bahamian shells are almost identical to the Northeastern Atlantic and Mediterranean ones in overall shape, arched in profile and with similar concentric growth lines. The protoconch is also paucispiral, consisting of 1.75 smooth whorls, but somewhat smaller in total diameter than the eastern shells (250 um vs. approximately 330 um, Valdéz & Lozouet, 2000; Silva & Landau, 2007). The most striking difference between the Bahamian and the eastern specimens lies in the position of the apex. In the Bahamian specimens the apex lies considerably more marginally; more posterior and further to the left than in the eastern shells. An important morphological feature of the eastern shells is a narrow rectilinear sulcus running obliquely from the apex to the edge, absent in the Bahamian shells. Discussion: The presence of these Spiricella shells in the Bahamas posed the question of whether one or two Recent species exist. From a morphological point of view, despite the lack of characters, there are two consistent differences between the eastern and western shells: the position of the apex and the presence or absence of the sulcus. Although the eastern specimens show some variability in the position of the apex, none comes close to that seen in the Bahamian shells (Figure 10). All the Northeastern Atlantic and Mediterranean shells show a more or less well developed sulcus associated with the apex, which is absent in the two Bahamian specimens. Apart from the differences outlined above, the overall diameter of the protoconch of the Bahamian shells is smaller than that of the eastern specimens. We therefore consider the Caribbean shells to represent a second species of Spiricella, S. redferni n. sp. We are not aware of any other fossil or Recent record for the genus in the Americas, or anywhere else outside Western Europe, the Mediterranean and Northwestern Africa. The European literature for the Page 308 genus, however, extends back to the Lower Oligocene, Rupelian Stage, of France (Waldéz & Lozouet, 2000). The fossil record would therefore suggest that Spiricella originated in the Eastern Atlantic and _ dispersed westwards. Interestingly, Vermeij & Rosenberg (1993) noted that many of the taxa in their list of westward- dispersing species had no fossil record in the Western Atlantic and appeared to be relatively recent immi- grants to the American coasts, but unlike many other westward invaders Spiricella apparently has not achieved a wide distribution in the Americas. Acknowledgments. We would like to thank Mr. Colin Redfern for bringing these interesting specimens to our notice, and donating the type material. REFERENCES CARROZZA, F. & R. ROCCHINI. 1987. Spiricella unguiculus Rang, 1827 (Gastropoda, Euthyneura: Umbraculidae) in the Mediterranean. Basteria 51:63—65. GeEUZE, G. J. & D. F. HOEKSEMA. 1994. New records of Spiricella unguiculus Rang, 1827 (Gastropoda Euthy- neura: Umbraculidae). Basteria 58:225—228. HOEKSEMA, D. F. & A. W. JANSSEN. 1984. Rediscovery of the marine gastropod Spiricella unguiculus Rang, 1827 (Eu- The Veliger, Vol. 50, No. 4 thyneura, Umbraculidae) in Miocene deposits of the North Sea Basin and in the Recent fauna of South West Europe. Basteria 48:7-11. JANSSEN, A. W. 1984. Mollusken uit het Mioceen van Winterswijk-Mist. Een inventarisatie met beschrijvingen en afbeeldingen van alle aangetroffen soorten. Konink- lijke Nederlandse Natuurhistorische Vereniging. Neder- landse Geologische Vereniging & Rijksmuseum van Geologie en Mineralogie: Amsterdam. 451 pp. RANG, P. C. & C. DES MOULINS. 1828. Description de trois genres nouveaux de coquille fossile du terrain tertiaire de Bordeaux, savoir: Spiricella, par M. Rang, correspondant; Gratelupia et Jouannetia, par M. Charles des Moulins, président. Bulletin d’Histoire Naturelle de la Societé Linéenne de Bordeaux 12:226—255. SILVA, C. M. & B. M. LANDAU. [2006] 2007. On the presence of Spiricella unguiculus Rang & Des Moulins, 1828 (Gastropoda, Notaspidea) in the European Pliocene: filling the geological gap. The Veliger 49(1):19-26. VALDES, A. & P. LOZOUET. 2000. Opisthobranch Molluscs from the Tertiary of the Aquitaine basin (southwestern France), with descriptions of seven new species and a new genus. Palaeontology 43(3):457-459. VERMEW, G. J. & G. ROSENBERG. 1993. Giving and receiving: the tropical Atlantic as donor and recipient region for invading species. American Malacological Bulletin 10: 181-194. THE VELIGER > 9) The Veliger 50(4):309-325 (December 16, 2008) © CMS, Inc., 2007 Functional Anatomy of Bankia fimbriatula Moll & Roch, 1931 (Bivalvia: Teredinidae) MARIA JULIA MARTINS-SILVA! AND WALTER NARCHI' "Departamento de Zoologia da Universidade de Brasilia, Campus Universitario, Asa Norte, 70910-900 Brasilia, DF, Brazil (e-mail: myjsilva@unb.br) Abstract. Bankia fimbriatula Moll & Roch, 1931, is a highly specialized bivalve adapted for boring into wood. Specimens were collected alive from a mangrove region at Praia Dura, Ubatuba, Sao Paulo, Brazil and maintained in an aquarium at room temperature (21°C) at the laboratory of the Department of Zoology, University of Sao Paulo. Studies of the anatomy were carried out on both relaxed and preserved specimens. Special attention was paid to the siphons, pallets, ctenidia, labial palps and mantle. The siphons are fairly long, and separated. The inhalant and exhalant siphons have ciliary activity at the tentacles, as described previously for Nausitora fusticula (Jeffreys, 1860). The posterior ctenidia are homorhabdic. Each ctenidium of B. fimbriatula is formed by the external demibranch only, with the blades in a “V” form. The ctenidia, associated with the rejection tracts of the mantle, present a good mechanism to deal with large quantities of particles, probably an adaptation for life in turbid waters. The labial palps are extremely reduced. The functional anatomy of B. fimbriatula suggests that both plankton and wood probably are important as food for this species. INTRODUCTION The anatomy of species of Teredinidae has been studied by several authors, including Quatrefages (1849), Menegaux (1889), Ridewood (1903), Sigerfoos (1908), Potts (1923), Lazier (1924), Atkins (1937), Purchon (1939, 1941, 1960), Nair (1957), Bade et al. (1961, 1964a, 1964b), Turner (1966), Rancurel (1971), Sar- aswathy & Nair (1971), Lopes & Narchi (1998) and Lopes et al. (1998). Some references [Atkins (1937), Purchon (1941, 1960), Clapp (1951), Morton (1970, 1978), Martinez (1987), Lopes & Narchi (1998) and Lopes et al. (1998), are restricted mainly to the structure and function of specific organs. The functional anatomy of Bankia fimbriatula Moll & Roch, 1931, is the main focus of this work; we analyzed the functioning of the siphons, the muscles associated with the siphons and pallets, and the ciliary currents related to the selection of food and particles for elimination. A detailed study of the anatomy and function of the stomach will be presented in a separate paper. Bankia fimbriatula occurs mainly in tropical warm waters around the world (Turner, 1966). On the Brazilian coast it has been reported from the littoral zone of Sao Paulo State (Lopes & Narchi, 1998), Parana State (Muller & Lana, 1986, 1987; Skinner et al., 1993) and Rio de Janeiro State (Silva et al., 1989; Junqueira et al., 1989; Martins-Silva et al., 1990 and Junqueira & Silva, 1991). 'Dr Walter Narchi (died in 2004) The genus Bankia Gray, 1842, includes twenty-three species (Turner, 1971) in the subfamily Bankiinae Turner, 1966, thirteen of which occur in the Brazilian littoral. Among species of Bankia, Sigerfoos (1908) studied the anatomy of Xylotrya gouldi [= Bankia gouldi (Bartsch, 1908)], Clapp (1951) made observations on living Teredinidae and described the siphons of B. gouldi. Bade et al. (1961) illustrated and described the mantle of Bankia minima [= Bankia carinata (Gray, 1827); Turner (1966, 1971)]. Bade et al. (1964a, 1964b) studied the digestive and respiratory systems of the same species. Turner (1966) described the posterior region of B. gouldi, and described the anatomy of Bankia setacea (Tryon, 1863), B. campanellata Roch & Moll, 1931, and B. australis (Calman, 1920). Sarasw- athy & Nair (1971) described the anatomy of B. indica. Tan et al. (1993) made a study of the shell and pallets of the early developmental stages of B. gouldi. This paper presents the first anatomical study of Bankia fimbriatula. MATERIAL AND METHODS Specimens of B. fimbriatula were collected during 1992 and 1994 in mangrove trees at Praia Dura, Ubatuba, Sao Paulo, Brazil (45°15’W, 23°30’S) (Figure 1). This is the second most abundant species of Teredinidae in the area, living at a salinity range from 0—33%o0 (Lopes & Narchi, 1993). The animals were kept inside the wood, in a seawater aquarium with constant aeration and at a salinity of 20%o and a room temperature of 22°C, where Page 310 _Z7 HIGHWAY The Veliger, Vol. 50, No. 4 MANGROVE 23° 30'S ATLANTIC OCEAN 160 240m —EE EEE Figure 1. Map of the mangrove region at Praia Dura, Ubatuba, Sao Paulo, Brazil, with indications of the collection stations (*1 and 2). they stayed in good condition for almost two years. The study of the functional anatomy was conducted at the University of Brasilia where the animals were kept alive in an aquarium with artificial seawater. Around 50 living and preserved specimens specimens of all sizes were analyzed. Identification of the material was based on Clench & Turner (1946) and Turner (1966, 1971). Identification was confirmed by the late Dr. Ruth D. Turner, Harvard University, USA. A lot of 15 complete specimens (shell, pallets and soft parts) were deposited in the Museu de Zoologia, Universidade de Sao Paulo (MZ USP) under the registration number 32061. Ciliary currents were studied by the application of a suspension of Carborundum (F3), carmine and Aqua- dag solutions. To help the observation of the different organs, whole animals were stained with Paracarmin and later cleared (Bicherl, 1943). Some of the anatomical details analyzed were also obtained by transverse sections 6-8 um thick, of M. J. Martins-Silva & W. Narchi, 2007 Page 311 PS UA) reef SUT Dust 0,5¢m Figure 2. Bankia fimbriatula. General topography of the organs after removal of the valve and the mantle from the left side of the body. Abbreviations: a, appendix; aa, anterior adductor muscle; ac, anterior ctenidium; an, anus; as, anterior stomach; au, auricle; bu, bulblike swelling; ch, cephalic hood; dh, dorsal hood; dd;, dds, digestive diverticles; ec, epibranchial cavity; ebv, efferent branchial duct; ex, exhalant siphon; f, foot; go, gonad; 1, intestine; in, inhalant siphon; k, kidney; m, mantle; ms, meddian stomach; p, pallet; pa, posterior adductor muscle; pc, posterior ctenidium; spc, semi-spiral conical projection; vc, ventricle. animals fixed in Bouin’s fluid and stained with Mallory’s Triple Stain or Ehrlichs’s haematoxylin and eosin, according to the method described by Pantin (1948). RESULTS General disposition of organs in the mantle cavity The disposition of the major organs in the mantle cavity of B. fimbriatula is shown in Figure 2. The visceral mass occupies about 60% of the body length and the posterior ctenidia and siphons occupy the remainder. The stomach has two regions; the appendix or wood- storing caecum is well developed and in the live animal is easily distinguished from the other structures because of its reddish color due to particles inside it. The digestive diverticula are of two types as defined by Potts (1923) and Morton (1970) for Teredo navalis (Linnaeus, 1758): the normal type and the specialized type. In live specimens of B. fimbriatula there is no difference between the digestive diverticula in contrast to what Lopes & Narchi (1998) described for Nausitora fusticula (Jeffreys, 1860). In B. fimbriatula the differ- ences are only distinct in histological sections. Males and females have milky white gonads, situated at the region immediately posterior to the distal part of the digestive diverticula. Contrary to what Turner (1966) states for B. gouldi, B setacea, B. campanellata and B. australis, the heart of B. fimbriatula has two intensely dark brown pigmen- tated atria. The ventricle is whitish in color and from it arises a well-developed aorta located on the dorsal surface of the gonads. The kidney is dorsal to the aorta, extending from the posterior part of the posterior adductor muscle to the distal extremity of the pericardial cavity. The nephro- stome opens into the interior of the pericardial cavity and the nephridiopores into the epibranchial cavity, both at the same level of the body. The afferent excretory duct shows, just behind the nephrostome, a globular dilatation whose internal wall is deeply folded and ciliated. The two nephridiopores are placed near each other, being smaller than, and situated posteriorly to, the gonopores. The anal canal lies in the dorsal region of the visceral mass and extends from the anus to the posterior end of Page 312 the gonads, connecting to the epibranchial cavity through a narrow opening. Shell The descriptive terminology of the shell is based on Turner (1966). Less than half the external surface of the shell of B. fimbriatula (Figure 3) is occupied by the anterior slope. The dorsal region possesses denticulate ridges that are eroded by friction against wood; laterally these ridges are more developed. The umbo- nal-ventral sulcus is narrow and flat. The dorsal and ventral condyles are obvious, but the umbonal-ventral ridge is poorly defined. The apophysis is flat when viewed transversely, with a sharp extremity near the ventral condyle. The posterior adductor muscle scar is only weakly evident. Pallets The pallets of B. fimbriatula (Figure 4) possess a long stalk, of the same length as or shorter than the blade. The pallets are elongate and the blades are composed of numerous cone-like elements on a central stalk; these elements are separated and easily removed from the stalk, particularly in dried specimens. The cones have a calcareous base covered with periostracum, which extends as a border. The width and the ornamentation of the periostracal border vary greatly; the border may be smooth, coarsely to finely serrate, or produced laterally as awns. Variations in the form of the blade could not be related to the age of the animals or to environmental conditions. As all specimens came from the same population and similar ecological conditions this can be interpreted as individual variation, as Lopes & Narchi (1998) observed for Nausitora fusticula. Siphons As described for WN. fusticula the inhalant and exhalant siphons (Figure 5) are joined for almost half of their length (Lopes & Narchi, 1998). Most of the specimens have white siphons with small spots of reddish brown pigmentation from the region of separation of the siphons to the aperture. This pigmentation is more abundant on the ventral sides of the inhalant siphon and on the dorsal side of the exhalant. The inhalant siphon (Figure 6) is fringed with a row of eight digitiform tentacles, between which the epithelium forms simple projections. When the animal is pumping water, the siphons project through the wood, and the digitiform tentacles are kept almost perpendicular to the axis of the siphons. The exhalant siphon possesses a relatively narrow The Veliger, Vol. 50, No. 4 opening whereas the margin is smooth and _ lacks digitiform projections. The exhalant siphon stretches and moves more actively than the inhalant, the latter stayed in the same position for a long period, moving only when disturbed or in order to quickly close the opening by flexing the digitiform projections. This movement apparently occurred without tactile stimulus and was not regular. The cilia on the siphons (Figure 7) occur mainly on the digitiform projections and the epithelium at the inhalant and the extremity of the exhalant siphons and _ produce a weak rejecting current that transports small particles outwards. Thus, these cilia contribute to cleaning the tentacle surface, impeding the settling of small particles. Bankia fimbriatula eliminates fecal pellets and pseudofeces via small jets. The exhalant and inhalant siphons, respectively, eject them a short distance away from the opening in the wood, where generally they accumulate. In the aquarium, large quantities of this waste accumulate, requiring weekly removal. Musculature of the pallets and siphons The musculature involved in moving the pallets and siphons (Figure 8) was described by Turner (1966) for Bankia gouldi and B. setacea. In B. fimbriatula, the musculature is similar to that of B. gouldi and includes the protractors, anterior retractors, median and poste- rior retractors and adductor muscles. These muscles unique to the Teredinidae are fixed to the proximal third part of the pallet stalk. The protactor muscle (pmp) of each pallet is composed of two well-developed bundles, easily seen externally as an open fan shape, with the narrower part directed to the anterior region of the animal. The muscle itself is fixed to the stalk and to the calcareous part of the gallery wall. The anterior retractor muscle (armp) of pallet is formed by two muscular bundles, the thicker “‘internal” and the thinner “‘external.” The internal is fixed to the internal face of the stalk and the external is fixed to the external face of the stalk. The posterior retractor muscle (prmp) is slim with little branching. The posterior retractor muscle ends inside the mantle and is not attached to any hard structure. The adductor muscle of the pallets (amp), the extremities of which are fixed to the internal face of the stalk, bring together the two pallets. In the body region where the musculature of the pallets occurs, it is possible to observe two well developed cylindrical muscular bundles of the retractor muscles of the siphon (rms). When the animal is pumping water, the pallets remain inside the gallery. Any disturbance in the environment causes retraction of the siphons. At this M. J. Martins-Silva & W. Narchi, 2007 Page 313 A 2mm B Figure 3. Bankia fimbriatula. Right valve of the shell. A. External view. B. Internal view. Abbreviations: ap, apophysis; as, anterior slope; aur, auricle; c, chondrophore; d, disc; dc, dorsal condyle; ps, posterior slope; vc, ventral condyle; vu, ventral umbonal sulcus. Page 314 The Veliger, Vol. 50, No. 4 ii anv ie ~ vith H Y A Bey) tim oF “| Si 1) aera Mee) | hy ; [Mi iii i; ts Los Figure 4. Bankia fimbriatula. Variations in the pallet. A. Pallet with complete peduncle. B. External view of the pallet. C. Internal view of the pallet. Abbreviations: aw, awns; bl, blade; co, cone; sk, stalk. time, the pallets are pushed into the opening of the gallery by contraction of the pallet protractor muscles. When the disturbance ceases, the pallets retract and the siphons extend out to the exterior. Pallet retraction is executed by retractor muscles at the same time that the adductors contract, thus moving the pallets’ blades apart, allowing for passage of the siphons. During this process, the protractor muscle of the pallet and the retractors of the siphons remain relaxed. Mantle The structure of the mantle is similar throughout the family (Turner, 1966). In B. fimbriatula the mantle is thin and transparent in the anterior third of the body. At the median third it is a little thicker, while at the posterior third, it is very thick. The tissue of the mantle is filled with a whitish substance. Groups of round granules of a reddish-brown color, as described by Lopes & Narchi (1998) in N. fusticula, are not present in B. fimbriatula. In the hypobranchial cavity at each side of the body, the internal epithelium of the mantle has a tract with well-developed cilia that extends from the anterior region to the base of the inhalant siphon. In the anterior and median regions of the body, these tracts are lateral. At the beginning of the posterior ctenidia they approach one another, meeting and becoming ventral. The mantle in the epibranchial cavity dorsal to the posterior ctenidia, near the siphons, has internally a thick zone of mucus cells, as also described by Nair (1957) and Saraswathy & Nair (1971) for B. carinata. Labial palps The labial palps of B. fimbriatula (Figure 9) are attached to the epithelium of the visceral mass (Turner, 1966). They are inconspicuous, the external and the M. J. Martins-Silva & W. Narchi, 2007 Page 315 Figure 5. Bankia fimbriatula. Siphons and pallets as observed in living animal removed from the wood. Abbreviations: fp, fecal pellet; in, inhalant siphon; ex, exhalant siphon; p, pallet. The arrows show the direction of the inhalant and exhalant currents. internal ones occupying, respectively, dorsal and a ventral positions. The dorsal palp is reduced to two flat folds. The ventral palp is reduced to a small, long and narrow elevation extending from the ventral border of the mouth to the anterior extremity of the marginal groove. Identification of the palps was possible only for some specimens. The ciliary currents were observed only in a few specimens. Ciliary activity was slight and movements of the particles near the mouth were not detected. Page 316 B The Veliger, Vol. 50, No. 4 ku uu Figure 6. Bankia fimbriatula. Detail of the aperture of the inhalant siphon with eight digitiform tentacles. A. Siphon opened with extended tentacles. B. Siphon showing tentacles bending across the aperture. Ctenidia The terminology adopted for the description of the ctenidia of B. fimbriatula is the same used by Ridewood (1903); Atkins (1937); Purchon (1939) and Lopes & Narchi (1998). The anterior ctenidia have from eight to nine filaments that correspond to those of the external lamella of the demibranch (Figure 10). Each filament is reduced to a simple bar, joined throughout its length to the epithelium of the visceral mass. The first and the last filaments are really semi-filaments because there is complete ciliation on only one of the lateral faces, as Sigerfoos (1908) described for B. gouldi. Depending on the condition of the body contraction, the ctenidium may become strongly folded, simulating a plait (Figure 11). The body of B. fimbriatula can contract to half its length in preserved animals or even in live ones removed from wood. The posterior part of the body is more affected by this contraction and the ctenidia become shorter and folded. The posterior ctenidia of B. fimbriatula are similar to those described by Lopes & Narchi (1998) for N. fusticula. The posterior ctenidia are represented only by the external demibranch [Purchon (1939, 1941) and Lopes & Narchi (1998)]. The demibranchs of B. fimbriatula are eulamelli- branch and homorhabdic. Each demibranch has a V- shaped form; the apex possesses a marginal groove 95 um deep. Each filament of the posterior ctenidia measures 40.8 um in width along practically its entire length. The free extremity is slightly dilated. Each filament (Figures 12, 13) has two bands of frontal cilia (fc) laterally disposed, each of which measures around 6.8 um in length, bordered by two rows of lateral- M. J. Martins-Silva & W. Narchi, 2007 Page 317 O,} eam Figure 7. Bankia fimbriatula. Rim of the inhalant siphon showing one digitiform tentacle and one simple projection occurring between two tentacles. Abbreviations: c, cilia; tc, cilia tufts. frontal cilia (lfc) of around 50 um in length; laterally, between the base of the filament and the lateral-frontal cilia, there are lateral cilia (Ic) around 23.8 um in length forming on each face a strip about 20.4 um in width. The frontal cilia of the lateral regions of the free extremity of the filaments are the same length and cover the top of the filaments. In this region, there are no large cilia which could be identified as being terminal. The rows of lateral-frontal and frontal cilia end at the marginal groove base and are of similar length as the frontal cilia. The posterior demibranchs of B. fimbriatula, joined by their respective ctenidial axes, separate at the posterior region of the visceral mass. The filaments become progressively smaller until they are reduced to the marginal groove. This groove is situated laterally on the visceral mass it extends to the anterior ctenidia, and is bordered by ciliated cells. In the anterior and median regions of the body the ctenidia are reduced to a marginal groove. The quantity of material transported inside the marginal groove is generally small. Excess particles are conducted to the anterior region where they accumulate, surrounded by mucus, and formed into large masses, which flow from Page 318 The Veliger, Vol. 50, No. 4 Figure 8. Bankia fimbriatula. Diagrammatic view of the muscles associated to the pallets and siphons. Abbreviations: amp, adductor muscle of the pallet; ex, exhalant siphon; in, inhalant siphon; p, pallet; pc, posterior ctenidia; pmp, protractor muscles of the pallet; rms, retractor muscle of the siphon; rmp, retractor muscle of the pallets. Ud L Figure 9. Bankia fimbriatula. Frontal view of the anterior extremity. Abbreviations: ch, dorsal hood; dlp, dorsal labial palp; f, foot sole; mo, mouth; s, shell; vc, ventral condyle; vlp, ventral labial palp. M. J. Martins-Silva & W. Narchi, 2007 Page 319 inside the groove. These masses are then transported by the cilia of the rejection tract of the mantle and eliminated. Ciliary Currents On the filament of each demibranch of the posterior ctenidia, the lateral cilia produce strong water currents, which aid in respiration and feeding. On one face of the filament these cilia produce a ventrally-directed cur- rent, and on the other, a dorsally-directed one. The lateral-frontal cilia project toward the sides of the filaments and alternatively cross with the adjacent B Figure 10. Bankia fimbriatula. Topography of the anterior region of the body after removal of the valves and submitted the animal to a process of diaphanization. A. View from the right side. B. View from the left side. Abbreviations: ac, anterior ctenidia; Dgl, Deshayes glands; dh, dorsal hood; mg, marginal groove; sa3, sorting area sa3. filaments to form a type of grating. The lateral cilia beat from the inside out onto the interfilamentar spaces, throwing particles onto the frontal faces of the filament. From here the particles are transported by the two lateral rows of the frontal cilia and large particles are prevented from penetrating in the interior of the demibranchs. The frontal cilia on the external and internal blade of each demibranch conduct particles of different sizes to the ventral region. At the ventral extremity of the filaments, small particles are moved by the cilia from the lateral faces to the interior of the marginal groove and conducted to the Page 320 anterior region by a strong ciliary current. Larger particles are conducted anteriorly at the free edge of the demibranch and outside the marginal groove. Opening and closing of the marginal groove were not observed to control the quantities of particles within. Depending on the distance of the marginal groove from the mantle rejection tracts, the strong ciliary currents of these tracts directly captured particles from outside the The Veliger, Vol. 50, No. 4 3 © wot Oo Figure 11. Bankia fimbriatula. Posterior ctenidium. A. Diagrammatic vertical section through the ctenidium to show the mode of action of the frontal cilia. B. horizontal section to show the homorhabdic condition of the ctenidium. Abbreviations: alod, ascendent lamella of the outer demibranch; dlod, descendent lamella of the outer demibranch; ifj, interfilamentar junction; mg, marginal groove. marginal groove. Particles that form large masses that fall into the hypobranchial cavity are collected by the rejection tract. These masses are retained by the cilia of the rejection tract of the mantle and then eliminated. Weak ciliary activity was observed in the mid-region of the visceral mass in a few specimens of B. fimbriatula, particles being conducted forward in the marginal groove. M. J. Martins-Silva & W. Narchi, 2007 A < ‘ Paes Pas icp htt es Page 321 O = GE aE ETE 7 . Sf fi fj} La Figure 12. Bankia fimbriatula. Marginal groove at the ventral margin of the ctenidium. A. Cilia on the outer surface of the ctenidium; a—b, line of the marginal groove. B. Outer surface of a filament showing the cilia. C. Frontal view between two filaments to show cilia. Abbreviations: fc, frontal cilia; lc, lateral cilia; lfc, latero-frontal cilia; the arrows indicate the direction of ciliary currents, the arrows indicate direction of ciliary currents, including oral one. The material entering the epibranchial include feces, gametes, excretory products and very small particles which have passed through the demibranchs. Beside the ciliary currents on the epithelium, frequent contractions were observed throughout the length of the mantle at the epibranchial cavity. The feces were eliminated by short and intermittent jets. No ciliary activity was detected on the wall of the anal canal. DISCUSSION The eight simple digitiform tentacles of the inhalant siphon of B. fimbriatula and their projection within the inhalant aperture do not act as barriers against particles entering the mantle cavity, but the siphon can regulate the quantity of the material that enters by contracting the circular muscle at the base of the Page 322 25 um ate ss Perce The Veliger, Vol. 50, No. 4 Figure 13. | Bankia fimbriatula. Transverse sections of the filaments to show cilia. A. Apical filaments. B. Basal filaments. Abbreviations: ch, chitin; fc, frontal cilia; Ic, lateral cilia; lfc, latero-frontal cilia; m, horizontal muscles. tentacle or withdrawing into the interior of the gallery. The ciliary currents of the ctenidia and the efficient rejection tracts of the mantle are responsible for the elimination of rejected particles. Ciliary activity observed on the tentacles of the inhalant siphon is similar to that described for N. fusticula (Lopes & Narchi, 1998) and is apparently related to the removal of small particles which settle around and accumulate on these structures. The retraction and extension movements of the siphons are probably related to the cleaning mechanism, eliminating larger particles which are not removed by ciliary action. M. J. Martins-Silva & W. Narchi, 2007 The exhalant siphon is longer than the inhalant and has no digitiform tentacles at the exhalant opening. The exhalant siphon has great flexibility and shows movements of retraction and extension that can make it increase about three times in length. It has many cilia that remove small particles and it is probably sensitive to mechanical stimuli. The tentacles of the inhalant siphon were described by Townsley et al. (1965) for B. setacea and by Lopes & Narchi (1998) for N. fusticula. The tentacles of B. fimbriatula are different from those described by Lopes & Narchi (1998). The sensitivity of the siphons of B. fimbriatula to mechanical stimuli, even of low intensity, is similar to that reported for other teredinids (Quatrefages, 1849; Saraswathy & Nair, 1971; Lopes & Narchi, 1998). The siphons of B. fimbriatula, contrary to Turner (1966), are not fairly long and separate, but are united at the basal region for at least one third their length. The mantle of B. fimbriatula is similar to that of other teredinids (Sigerfoos, 1908; Lazier, 1924; Turner, 1966; Saraswathy & Nair, 1971; Rancurel, 1971 and Lopes & Narchi, 1998). A thick mantle is apparently generally distributed throughout the family, especially in older specimens, but is nowhere as great as described in Kuphus (Guettard, 1770; Turner, 1966). It has been described for Bankia by Bade et al. (1961), Sigerfoos (1908) mentions it for B. gouldi; Turner (1966) for Bactronophorus YVapparone-Canefri, 1877, Neoteredo Bartsch, 1920 and Nausitora Wright, 1864 and Lopes & Narchi (1998) for N. fusticula. In all species having a thick mantle there were also clusters of red-brown, berry-like structures on the transverse fibers of the middle layers. These structures are not present in B. fimbriatula. The rejection tracts of the mantle in B. fimbriatula are separate throughout their length as was observed by Sigerfoos (1908) in B. gouldi, by Saraswathy & Nair (1971) in Nausitora hedleyi Shepman, 1919 and by Lopes & Narchi (1998) in N. fusticula. In Teredo norvegica [= Nototeredo norvegica (Spengler, 1792)] and Teredo megotara [= Psiloteredo megotara (Hanley, 1848)], studied by Saraswathy & Nair (1971), these tracts are fused at the posterior body region. In B. fimbriatula, these tracts are fused at the posterior body region on the ventral side up to the basal region of the inhalant siphon. The anterior ctenidium of B. fimbriatula is composed of eight or nine filaments. In B. gou/di the number of filaments is usually nine, rarely 10 or 11 (Sigerfoos, 1908). Some authors noted a constant number of filaments: Purchon (1941) recorded ten for N. norvegica (= Nototeredo norvagica)and seven for P. megotara; Saraswathy & Nair (1971), eight for N. hedleyi and six for Teredo furcifera von Martens, 1894. According to Lazier (1924), there are five filaments in 7. navalis, Page 323 whereas according to Morton (1970), there are eight. Lopes & Narchi (1998) in N. fusticula recorded from six to eight filaments, depending on the specimen. This variation is not correlated with the size of the animal. According to Lopes & Narchi (1998) the absence of variation in filament number or the discrepancy in numbers of filaments in the same species could be related to small sample size. The posterior ctenidium of B. fimbriatula occupies more than 50% of the body length. According to Turner (1966) species with long ctenidia and more developed palps feed mainly on plankton, wood being a less important source to the animal. The appendix of B. fimbriatula makes up 1/3 of the body length, showing that wood is an important source of food. The species have a large appendix and the posterior ctenidia are also well developed. This suggests that plankton, as well as wood particles are important in the nutrition of this species. In B. fimbriatula, the basic anatomy of the ctenidia does not significantly differ from that of other teredinid species described by Ridewood (1903), Sigerfoos (1908) and Lopes & Narchi (1998). In B. fimbriatula, however, the fine frontal cilia are laterally disposed in two tracts in the filament, and the central part is free of cilia. Only at the terminal part of the filament, near the marginal groove, fine cilia cover the entire marginal tip of the filament. Comparing the anatomical characters described in the present work for B. fimbriatula with those of B. gouldi described by Sigerfoos (1908), Clapp (1951), Turner (1966) and Tan et al. (1993) we conclude that B. fimbriatula differs from B. gouldi in terms of the pallets, the presence of eight tentacles around the inhalant siphon and the deep marginal groove in the posterior ctenidium. Bankia fimbriatula differs also from B. gouldi, B. setacea, B. campanellata and B. australis by the presence of tubular auricles intensively pigmented (Turner, 1966). In addition, B. fimbriatula differs from B. carinata studied by Bade et al. (1961) and Sarasw- athy & Nair (1971) in the position of the gonads and the large size of the appendix and the great develop- ment of the posterior ctenidia. According to Turner (1966) the variation exhibited by dissected individuals is considerable. Within the range of the genus much more additional work will be necessary before the many subgenera described on the basis of the pallets can be evaluated. Acknowledgments. The authors wish to express their gratitude to the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico - CNPq - Brazil, for the award of grants that made the present work possible. We also thank Mariana Zatz and Pedro De Podesta Uchéa de Aquino for illustration of Figures | and 2. The senior author expresses her pleasure at having worked with Dr Walter Narchi, who passed away in 2004. He was an Page 324 excellent mollusc researcher and his papers, published in major international journals, made the study of bivalve anatomy easy to carry out. Thank you Dr Walter very much for being my mentor. REFERENCES ATKINS, D. 1937. On the ciliary mechanisms and interrela- tionships of Lamellibranchs. Part II : Sorting devices on the gills. Quarterly Journal of Microscopical Sciences N.S. 79:339-373. BADE, I. V., V. B. MASUREKAR & D. V. BAL. 1961. A General account of the wood borer Bankia (Bankiella) minima. Bly., Nach., Roch. Journal of the University of Bombay 29(3/5):49-61. BADE, I. V., V. B. MASUREKAR & D. V. BAL. 1964a. Digestive system of the wood borer Bankia ( Bankiella) minima. Bly., Nach., Roch. Journal of the University of Bombay 32(3/5):52—59. BADE, I. V., V. B. MASUREKAR & D. V. BAL. 1964b. Respiratory system of the wood borer Bankia ( Bankiella) minima. Bly., Nach., Roch. Journal of the University of Bombay 32(3/5):60—70. ; BUCHERL, W. 1943. Compéndio de Técnica Microscopica. Ed. Anchieta Ltda: Sao Paulo. 307 pp. CLAPP, W. F. 1951. Observation on living Teredinidae. Fourth Progress Report [Rept. n° 7550], Wiliam F. Clapp Laboratoires Inc., Duxbury, Mass. pp. 1-9. CLENCH, W. J. & R. D. TURNER. 1946. The genus Bankia in the western Atlantic. Johnsonia, Cambridge 2:1—28. HiROKI, K., R. M. V. LEONEL & S. G. B. C. LOPES. 1994. Reproductive events of Nausitora fusticula (Jeffreys, 1960) (Mollusca, Bivalvia, Teredinidae). Invertebrate Repro- duction and Development 26(3):247-250. JUNQUEIRA, A. O. R., S. H. G. SILVA & M. J. MARTINS- SILVA. 1989. Avaliagao da infestagao e diversidade de Teredinidae (Mollusca - Bivalvia) ao longo da costa do Estado do Rio de Janeiro, Brasil. Memorias do Instituto Oswaldo Cruz 84(IV):275—280. JUNQUEIRA, A. O. R. & S. H. G. SILVA. 1991. Estudo experimental dos Teredinidae Rafinesque, 1815 (Mollusca : Bivalvia) no estuario da Lagoa da Tijuca, Rio de Janeiro, RJ, Brasil. Revista Brasileira de Biologia 51(1):113-126. LAZIER, E. L. 1924. Morphology of the digestive tract of Teredo navalis. University of California Publications in Zoology 22(14):455-474. Lopes, S. G. B. C. & W. NARCHI. 1993. Levantamento e distribuigao das espécies de Teredinidae (Mollusca Bivalvia) no manguezal da Praia Dura, Ubatuba, Sao Paulo, Brasil. Boletim do Instituto Oceanografico, Sao Paulo 41(1/2):29-38. Lopes, S. G. B. C. & W. NARCHI. 1998. Functional anatomy of Nausitora fusticula (Jeffreys, 1860) (Bivalvia: Teredini- dae). The Veliger 41(3):274-288. Lopes, S. G. B. C., W. NARCHI & O. DOMANESCHI. 1998. Digestive tract and functional anatomy of the stomach of Nausitora fusticula (Jeffreys, 1860) (Bivalvia: Teredini- dae). The Veliger 41(4):351—365. MARTINEZ, J. C. 1987. Structure et fonctionnement de Pappareil digestif de Teredo navalis L. (Teredinidae : Bivalvia). Haliotis 16:197—207. MARTINS-SILVA, M. J., A. O. R. JUNQUEIRA & S. H. G. SILVA. 1990. Distribuigao dos organismos marinhos perfurantes de madeira, segundo um gradiente crescente The Veliger, Volls05 Nos: de salinidade, no Canal de Itajuru, Cabo Frio, Rio de Janeiro, Brasil. IN: Simposio sobre Ecossistemas da Costa Sul Sudeste Brasileira 2:264-272. MENEGAUX, A. 1889. Sur les homologies de différents organes du Taret. Bulletin de la Societé du Zoologie Frangaise 14: 53-55. Morton, B. 1970. The functional anatomy of the organs of feeding and digestion of Teredo navalis Linnaeus and Lyrodus pedicellatus (Quatrefages, 1849). Proceedings of the Malacological Society of London 39(151):151—167. Morton, B. 1978. Feeding and digestion in shipworms. Oceanography and Marine Biology, Annual Review 16: 107-144. MULLER, A. C. P. & P. C. LANA. 1986. Teredinidae (Mollusca: Bivalvia) do litoral do Parana. Neritica, Pontal do Sul 1(3):27-48. MULLER, A. C. P. & P. C. LANA. 1987. Padrodes de distribuigao geografica de Teredinidae (Bivalvia : Mol- lusca) no Estado do Parana. Ciéncia e Cultura, Sao Paulo 39:1175-1177. Nair, N. B. 1957. Physiology of digestion in Bankia indica, the enzymatic activity of the crystalline style. Journal of Scientific Industrial Research 16C(2):39-41. PANTIN, C. F. A. 1948. Notes on microscopical technique for zoologists. University Press: Cambridge. 77 pp. Potts, F. A. 1923. The structure and function of the liver of Teredo, the shipworm. Proceedings of the Cambridge Philosophical Society, Biological Sciences 1(1):1—17. PURCHON, R. D. 1939. Reduction of ctenidia in the Lamellibranchia. Nature 144:206. PURCHON, R. D. 1941. On the biology and relationship f the Lamellibranch Xy/lotrya dorsalis (Turton). Journal of Marine Biological Association of the United Kingdom 25:1-39. PURCHON, R. D. 1960. The stomach in the Eulamellibanchia : stomach types IV and V. Proceedings of the Zoological Society of London 135(3):431-489. QUATREFAGES, A. 1849. Mémoire sur le genre Taret (Teredo Linn.). Annales des Sciences Naturelles, Zoologie, (Paris) 11(3):19-64. RANCUREL, P. 1971. Les Teredinidae (Mollusques lamelli- branches) dans les lagunes de Cote d’Ivoire. Mémoires Office de la Recherche Scientifique et Techinique Outre- mer, Paris 47:1—235. RIDEWOOD, W. G. 1903. On the structure of the gills of the Eulamellibranchia. Philosophical Transactions of the Royal Society of London 195B:147—284. SARASWATHY, M. & N. B. NAIR. 1971. Observations on the structure of the shipworms Nausitora headleyi, Teredo furcifera and Teredoa princesae (Bivalvia : Teredinidae). Transactions of the Royal Society of Edinburg 68(14): 507-566. SIGERFOOS, C. P. 1908. Natural history, organization and late development of the Teredinidae or shipworms. Bulletin of the Bureau of Fisheries, Washington 39:191—231. SILVA, S. H. G., A. O. R. JUNQUEIRA, M. J. MARTINS-SILVA, I. R. ZALMON & H. P. LAVRADO. 1989. Fouling and wood boring communities distribution on the coast of Rio de Janeiro, Brazil. Pp. 95-109. in C. Neves & O. T. Magoon (eds.), Coastline of Brazil. American Society of Civil Engineers: Charleston. SKINNER, L. F., S. H. G. SILVA & M. J. MARTINS-SILVA. 1993. Estudo das comunidades incrustantes e perfurantes ao longo do Canal do Bacalhau, Guaratiba/RJ Anais do M. J. Martins-Silva & W. Narchi, 2007 IIT Simposio sobre Ecossistemas da costa sul sudeste brasileira: 228-235. Tan, A. S., Y. Hu, M. CAsTaGna, R. A. Lutz, M. J. C. KENNISH & A. S. POOLEY. 1993. Shell and pallet morphology of early developmental stage of Bankia gouldi (Bartsch, 1908) (Bivalvia: Teredinidae). The Nautilus 107(2):63—75. TOWNSLEY, P. M., R. A. RICHy & P. C. RUSSE. 1965. The occurence of protoporphyrin and myoglobin in the marine borer Bankia setacea (Tryon). Canadian Journal of Zoology 43:167-172. TURNER, R. D. 1966. A survey and illustrated catalogue of the Paser3s25 Teredinidae (Mollusca: Bivalvia). The Museum of Com- parative Zoology, Harvard University: Cambridge. 265 pp. TURNER, R. D. 1971. Identification of marine- boring molluscs. Pp. 17-64 in E. B. G. Jones & S. K. Eltringham (eds.), Marine borers, fungi and fouling organisms of wood. Organization for Economic Co-Operation and Development: Paris. YONGE, C. M. 1957. Mantle fusion in the Lamellibranchia. Pubblicazioni della Stazioni Zoologica di Napoli 29:151— 171. The Veliger 50(4):326—328 (December 16, 2008) THE VELIGER © CMS, Inc., 2007 Drilling Localization on Bivalve Prey by Octopus rubescens Berry, 1953 (Cephalopoda: Octopodidae) ROLAND C. ANDERSON,' DAVID L. SINN? AND JENNIFER A. MATHER? 'The Seattle Aquarium, 1483 Alaskan Way, Seattle, WA USA (e-mail: roland.anderson@seattle.gov) "University of Tasmania, School of Zoology, Hobart, Australia *University of Lethbridge, Department of Psychology, Lethbridge, Canada One Sentence Abstract. An examination of 171 shells of clams (Venerupis philippinarum (Adams & Reeve, 1850)) eaten by Octopus rubescens Berry, 1953 showed that holes in them were drilled by the octopuses preferentially (64.3%) in adductor muscle scar areas (anterior or posterior), which together comprised only 6% of the total shell area. Key Words: octopus, drill holes, bivalves, adductor muscles. Octopuses are well-known generalist predators (Han- lon & Messenger, 1996), but within this generalist approach they also display individual dietary prefer- ences (Anderson & Mather, 2007) and feeding methods (Dodge & Scheel, 1999). Bivalves make up a substantial part of the diet of many octopuses and the methods octopuses use when drilling them, while time-consum- ing (Steer & Semmens, 2003), are not well-documented and appear to be highly variable (Anderson & Mather, 2007). After drilling, octopuses inject venom into clam prey in order to paralyze the muscle (Nixon & Maconnachie, 1988). Such energy expenditure in drilling might be minimized by selection of particular locations on the bivalve shell (Steer & Semmens, 2003). Anderson & Mather (2007) reported that Enteroctopus dofleini drills clams in the center of the shell. This is unlike O. vulgaris, which drills around the edge (Ambrose & Nelson, 1983), and O. dierythraeus Norman, 1993, O. mimus Gould, 1852 or O. bimacu- loides Pickford & MacConnaughey, 1949, which drill over the adductor muscles (Steer & Semmens, 2003; Cortez, et al., 1998; Casey, 1999, respectively). This inter-specific variation in drilling behavior highhghts the fact that one of the central problems octopuses face when feeding on bivalves, in addition to choice of prey, is where to drill on a clam’s shell, as different areas of the shell may be thicker or thinner and vital organs of the clam are located in species- specific areas (Kozloff, 1990). Observations of clams eaten by O. rubescens at the Seattle Aquarium indicated that individuals may learn to drill clams in particular locations (Anderson, et al., in prep.), while drilling efficiency appears to deteriorate during senescence (Anderson et al., 2008). Despite these observations on potential life-stage specific differences, there are no detailed studies of the localization of drill holes by mature O. rubescens, and that is the subject of this report. Ten Octopus rubescens (mean weight: 73.2 g; SD = 64.6) caught in the wild were held at the Seattle Aquarium and fed only Manila clams (Venerupis philippinarum, (Adams & Reeve, 1850)) obtained from a local fish market. At least ten shells from clams that had been drilled and eaten were then collected from each octopus over a period of a month (n = 171; an additional 187 were eaten but undrilled). All drilled shells had one hole in them. The holes were typically 1.4 mm wide on the surface (n = 30; SD = 0.28) and 0.4 mm wide on the inner surface of the shell (n = 30; SD = 0.15) as measured with a light microscope. The dimensions of the eaten shells and their adductor muscle scars were also measured and their areas calculated (nt X L * W/2). Locations of drill holes in shells were classified as occurring in the umbo, center, posterior, anterior, or ventral regions of a shell, by the methods of Anderson & Mather (2007) (see Figure 1) and further, whether they occurred within an adductor muscle scar. The mean shell length was 36.2 mm (SD = 4.57). The mean area of the anterior adductor muscle was 2.6% of the shell area and the posterior muscle scar was 3.7% (n = 171). We used a replicated test of goodness-of-fit (Sokal & Rohlf, 1995) to determine whether proportions of drill hole location (umbo, center, posterior, anterior, or ventral) differed significantly from 20:20:20:20:20. A significant result in the first analysis would indicate non-random targeting of particular areas of the shells. We again used a replicated goodness-of-fit test to R. C. Anderson et al., 2007 Umbo ‘a Anterior Adductor Muscle Scar Page 327 iB Center Posterior Adductor Muscle Scar \_Ventral Figure 1. Typical clam shell (Venerupis philippinarum) drilled by Octopus rubescens. The mean shell length was 36.2 mm. The mean area of the anterior adductor muscle was 2.6% that of the whole and the posterior muscle scar was 3.7% (n = 171). The areas are stylized but are roughly equal in area. determine whether drill hole location (over adductor muscles or outside adductor muscles) differed from the expected frequency of 6:94. A significant result in the second analyses would indicate that octopuses were actively targeting adductor muscles. Since octopuses could contribute to more than one observation in both analyses we first tested whether the outcomes of all the octopuses were homogeneous (i.e., heterogeneity G- test), that is, were individuals uniform with respect to frequencies of drill holes in the different regions of shell. After taking this octopus individuality into account, we then tested whether the sample as a whole fit the expected ratio of frequencies (i.e., results were pooled within each octopus: total G test). This approach allowed us to examine both individual-level as well as overall average pattern of drilling localization. Two of 10 individual octopuses drilled with equal probability in each of the five valve locations (hetero- geneity G-test = 100.85, df = 36, P < 0.05) but overall, there was still a clear significant preference for octopuses to drill in anterior regions of the clams (total G-test = 213.35, df = 40, P < 0.05). It was also clear that octopuses were targeting the adductor muscle scars: 64.3% of drill holes were in adductor muscle scars (anterior or posterior), which together comprised only 6% of the shell area. Once again, while some individuals did not drill over muscles as frequently as others (heterogeneity G-test = 40.15, df = 8, P < 0.05), there was still a strong significant overall trend for octopuses to drill within muscle scar areas (total G-test = 445.78, df = 10, P < 0.001). Although there are slight differences between exter- nal features of the anterior and posterior ends of Venerupis philippinarum (e.g., the anterior end is very slightly pointed and the posterior end is rounded, see: Coan et al., 2000), it is interesting to note that the majority of octopus drill holes were located in the anterior end (52% of all observed drill holes, 20% expected by chance alone) and that most individuals appeared to target the adductor muscles. Octopus rubescens is known for its potent venom (Hanlon & Messenger, 1996) so targeting adductor muscles which hold the clam shells closed (Kozloff, 1990) for venom injection and paralyzing one of the adductor muscles Table 1 Frequencies of drill hole locations found in different regions of clam shells left after predation by Octopus rubescens (n 171). Anterior Posterior Umbo Center Ventral Within muscle scar Total N 89 24 34 18 > 110 Percentage 52.0 14.0 20.5 10.5 DS) 64.3 Page 328 may be the most efficient means of accessing food. Cortez et al. (1998) hypothesize there may a direct effect on the nervous system of the clam by injecting venom in any anterior region of the clam. This brings up the interesting question of what features (physical and/or chemical) of clam shells octopuses use to gather information regarding internal location of clam organs and musculature. Given that half of the clams eaten during our study were not drilled at all, are these same cues used to determine whether to drill at all? Clearly, further studies are needed to ascertain the conditions which favor non-random drilling behavior in octopuses, including the apparent efficiency of octopuses at drilling shells from clam species with short co-existence histories and the maintenance of behavioral individu- ality and foraging strategies witnessed here and in other studies (Mather & Anderson, 1993; Sinn et al., 2001). Acknowledgments. We thank Richard Peters and Paul Black- low for helpful suggestions regarding data analysis and Barry Shuman prepared the figure. Greg Diet] and two anonymous reviewers provided constructive comments to earlier drafts of the manuscript. REFERENCES AMBROSE, R. F. & B. NELSON. 1983. Predation by Octopus vulgaris in the Mediterranean. Pubblicationi della Sta- zione Zoologica di Napoli 1: Marine Ecology 4:251—261. ANDERSON, R. C. & J. A. MATHER. 2007. The packaging problem: bivalve prey selection and prey entry techniques of the octopus Enteroctopus dofleini. Journal of Compar- ative Psychology 121:300—305. The Veliger, Vol. 50, No. 4 ANDERSON, R. C., J. A. MATHER & D. L. SINN. 2008. Octopus senescence: forgetting how to eat clams. The Festivus 40:55—56. CASEY, E. 1999. Intelligent predation by the California two- spot octopus. The Festivus 31:21—24. COAN, E. V., P. VALENTICH-SCOTT. & F. R. BERNARD. 2000. Bivalve seashells of western North America. Santa Barbara Museum of Natural History: Santa Barbara, CA. CorTEZ, T., B. G. CASTRO & A. GUERRA. 1995. Feeding dynamics of Octopus mimus (Mollusca: Cephalopoda) in northern Chile waters. Marine Biology 123:497—503. DopGE, R. & D. SCHEEL. 1999. Remains of the prey- recognizing the midden piles of Octopus dofleini (Wilker). The Veliger 42:260-266. HANLON, R. T. & J. B. MESSENGER. 1996. Cephalopod behaviour. Cambridge University Press: Cambridge. KOZLOFF, E. N. 1990. Invertebrates. Saunders College Publishing: Philadelphia. MATHER, J. A. & R. C. ANDERSON. 1993. Personalities of octopuses (Octopus rubescens). Journal of Comparative Psychology 107:336—340. Nixon, M. & E. MACONNACHIE. 1988. Drilling by Octopus vulgaris (Mollusca: Cephalopoda) in the Mediterranean. Journal of Zoology, London 216:687—716. SInn, D. L., N. A. PERRIN, J. A. MATHER & R. C. ANDERSON. 2001. Early temperamental traits in an octopus (Octopus bimaculoides). Journal of Comparative Psychology 115:351—364. SOKAL, R. R. & F. J. ROHLF. 1995. Biometry. W.H. Freeman and Company: New York. STEER, M. A. & J. M. SEMMENS. 2003. Pulling or drilling, does size or species matter? An experimental study of prey handling in Octopus dierythraeus (Norman, 1992). Journal of Marine Biology and Ecology 290:165—178. ; = ; 7 ca Si 1 7 ue ne i & ux tN ar Instructions to Authors The Veliger publishes original papers on any aspect of malacology. 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Send manuscripts, proofs, books for review, and correspondence on editorial matters to: Geerat J. Vermeij Editor, The Veliger Department of Geology University of California at Davis One Shields Avenue Davis, CA 95616 veliger@geology.ucdavis.edu TW 530:752.2234 F 530.752.0951 In the cover letter, authors should briefly state the point of the paper, and provide full and electronic addresses of at least three reviewers who have not previously seen the manuscript. If authors feel strongly that certain reviewers would be inappropriate, they should indicate reasons for their views. MITHSONIAN INSTITUTION Li WANN BRARIES Contents — Continued 3 9088 01435 tl RESEARCH NOTE Drilling Localization on Bivalve Prey by Octopus rubescens Berry, 1953 (Cephalopda: Octopo- didae) RoLaNp C. ANDERSON, Davin L. SINN AND JENNIFER A. 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