UE 4-0 | ISSN 0042-3211 We THE xwvz Pe ELIGE A Quarterly published by S24 } CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC." -S = f Berkeley, California we R. Stohler (1901-2000), Founding Editor Volume 51 March 31, 2010 Number 1 CONTENTS A Revised Classification of the Gastropod Genus Nerita INIETETSS AW Ate REN rset eee teeta Pane Se R Nr JU a Mea uN task mare vasa tap eldleal ave eign availa 1 Bostrycapulus heteropoma n. sp. and Bostrycapulus tegulicius (Gastropoda: Calyptraeidae) from Western Africa RAG HIET COLLINVAN DIE MITIOUROLAN Safe aoc co cis fetes iste tele cles wis cles v's sree ee a 8 A New Species of Hypselodoris and a Redescription of Hypselodoris picta lajensis (Nudibranchia: Chromodorididae) from Brazil Stone Dacosta, VINIcIuS PADULA AND MICHAEL SCHRODL...........0 000 cece eee 15 Diet and Feeding Habits of Octopus hubbsorum Berry, 1953, in the Central Mexican Pacific Ernesto L6pEz-UriartE, EDUARDO Rios-JARA AND MOnIca ELIZABETH (CONZATEZ ARO DRI CUE Zeenat cies oven sn See yiny a he Nok aula handed gs 26 Sacoglossan Opisthobranchs on Northwestern Pacific Shores: Stiliger berghi Baba, 1937, and Elysia sp. on Filamentous Red Algae Cyntuia D. TRowBrRIDGE, YosHIAKI J. HtRANO AND Yayor M. HirANO............--- 43 A New Species of Anatoma (Vetigastropoda: Anatomidae) from a Hydrothermal Vent Field in Myojin Knoll Caldera, Izu-Ogasawara Arc, Japan TAKENORI SasakI, DaNnteL L. GEIGER AND TAKASHI OKUTANI... 2-0-2000 0 eee eee eee ee 63 Oligocene and Miocene Vesicomyid Bivalves from the Katalla District, Southern Alaska STHPERENG GE VAIN ICA ZW IDAIKAWAINTAIN Oy sienna een tuas ai o es cyale a qs rai ti eae s 76 Trophonella (Gastropoda: Muricidae), a New Genus from Antarctic Waters, with the Description of a New Species MEG a TARASEWVCEVAND GUIDONPASTORINO 2.5 3c oa) sas ieee hse ose seh ee ee eee es 85 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. 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 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 David R. Lindberg, Museum of Paleontology, 1101 VLSB MC# 4780, University of California, Berkeley, CA 94720-4780, USA 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 E 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 Véliger. 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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: David R. Lindberg, Museum of Paleontology, 1101 VLSB MC# 4780, University of California, Berkeley, CA 94720-4780, USA. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). The Veliger 51(1):1-7 (March 31, 2010) Sire verierRe © CMS, Inc., 2008 A Revised Classification of the Gastropod Genus WNerita MELISSA A. FREY Center for Population Biology, University of California, Davis, Davis, California 95616, USA (e-mail: mafrey@ucdavis.edu) Abstract. The gastropod genus Nerita Linnaeus, 1758, is one of the most prominent groups found within tropical and subtropical marine intertidal communities. Yet until recently, a comprehensive systematic study of the group had not been undertaken. Using a newly reconstructed molecular phylogeny as a framework, I present a revised classification for the genus. I discuss the reassignment of several species, and review the molecular and morphological evidence in support of each subclade. INTRODUCTION The genus Nerita is a prominent group of marine intertidal gastropods that are widely distributed across tropical and subtropical shores. The clade ranges throughout each of the major oceans, with representa- tives inhabiting both continental coastlines and oceanic islands (Frey and Vermeij, 2008). Nerita grazes throughout the intertidal gradient, from the lower littoral zone to the supralittoral fringe, and occupies a variety of macrohabitats, including rocky intertidal, adjacent sands, mudflats, and mangroves. Given the group’s widespread distribution and abundance, spe- cies of Nerita have served in numerous ecological and evolutionary studies (e.g., Vermeij, 1973; Bertness and Cunningham, 1981; Garrity, 1984; Underwood, 1984; Underwood and Chapman, 1996; Waters et al., 2005; Hurtado et al., 2007; Crandall et al., 2008; Frey and Vermeij, 2008). Until recently, however, the systematic relationships among Nerita species remained poorly understood. Traditionally, the classification of Nerita species has been based on morphological characters. Since Lin- naeus (1758) first introduced the genus, authorities have divided the clade into multiple subgenera, adding new groups piecemeal with the recognition of shared traits (reviewed in Krijnen, 2002). The resulting subgenera have been defined largely by conchological and opercular features (Morch, 1852; Gray, 1858; Martens, 1887-1889; Vermeij, 1984; Dekker, 2000; Krijnen, 2002). However, variability within such traits and conflict among the characters has resulted in taxonomic confusion and unstable systematic group- ings. Radular characteristics, which exhibit little variation across the genus, have offered almost no phylogenetic utility (Komatsu, 1986), and other ana- tomical traits, such as reproductive structures, have been examined in only a few Nerita species (Fretter, 1965; Holthuis, 1995). Indeed, until recently, the only comprehensive, species-level treatment of Nerita fo- cused on nomenclature, conchological descriptions, and distributions (Krijnen et al., 1995; Krijnen et al., 1997; Krijnen et al., 1999; Krijnen et al., 2001, 2005). Yet, by delimiting morphospecies, these latter works have significantly improved Nerita taxonomy and have set the necessary groundwork for additional systematic studies. Advances in molecular systematics offer an oppor- tunity to build upon such knowledge and to develop a clear understanding of taxonomic relationships. Re- cently, a molecular phylogenetic study of Nerita was conducted in order to examine the group’s biogeo- graphic history (Frey and Vermeij, 2008). The study included representatives from each subgenus, and more than 85% of the species as currently defined, resulting in the most comprehensive systematic analysis of the group to date. Here, I use the resulting phylogeny as a framework to outline a new classification for the genus. A PHYLOGENETIC FRAMEWORK A species-level phylogeny of Nerita was previously reconstructed using multiple molecular markers and phylogenetic methods (Frey and Vermeij, 2008). From each species, both mitochondrial (16S and COI) and nuclear (ATPSa) genes were amplified and sequenced. Phylogenetic analyses, including Maximum Parsimony (MP), Maximum Likelihood (ML), and Bayesian (MB) methods, were conducted on each separate gene and on a combined dataset (for details, see Frey and Vermeij, 2008). Bootstrap proportions (=70%) and posterior probabilities (295%) were used to estimate clade support. A single tree, which yielded the highest- likelihood value recovered from a Bayesian analysis of the combined dataset, corresponded to the general topology of both the MP and MB consensus trees and showed minimal conflict with separate gene trees (Frey and Vermeij, 2008). Accordingly, this tree was selected Page 2 100/100/100 100/100/100 100/100/100 100/100/100 adenensis planospira lirellata atramentosa morio melanotragus 100/100/100 99/84/100 97/99100F" 00/99/100 . 00/96/10 100/100/100 100/100/100 94/82/- 100/100/100 98/88/99 100/99/100 100/99/100 87/96/98 100/100/100 100/94/99 -/80/99 100/100/100 100/-/ | 100/100/100 99/96/10 100/99/100 98/96/100 4100/1 100/ 100/100/100 100/100/100 | 194/100 10) -/-19B 99/91/100 1751 8 100/100/100 100/100/100 100/99/100_F"190/99/100 -/93/100 eS 100/100/100 100/97/100 100/100/100 86/70/- -/88/- -/87/- 100/100/100 100/100/100 100/100/100 99/72/- 400/100/100 196/100 97/10/99 80 98: Q0/100/10 99/100/100 86/93/100 100/100/100 98/99/100 -/95/100, 100/100/100 -/-198 100/100/100 100/97/100_F™5s;9n100 /900 99/96/10) poo 99/88/10 100/100/100 fiqo/se 1-198 83/81/100 1-196 100/90/100 100/96/100 82/-/95 10 100/100/101 100/100/100 == 100/400/101 96/98/98 =—"g 92/91/100 rmnncom esate 100/100/100 93/91/100 s00/100/100 | /74/- ri, 79/92/100 80/77/100 91/90/100 -/85/- MP BS/ML BS/MB PP 0.01 16S+COI+ATPSa Figure 1. and nuclear genes (Frey and Vermeij, 2008). Support for each clad funiculata ful gurans exuvia Japonica A Japonica B balt 100/300/100 scabricosta 10/1 Oteeeab 100/97/100 < 100/99/100 95/93/97 The Veliger, Vol. 51, No. 1 Adenerita Ilynerita Lisanerita tessellata senegalensis Theliostyla textilis 100/96/100 sanguinolenta -/84)- albicilla A albicilla B 100/89/100 100/92/100 Heminerita 00/100 longii — orbignyana polita polita B polita C antiquata = _Vitiensis erythrostoma litterata A litterata B 100/97/100 00/100/T00 100/100/100 10/95/1400 Linnerita 100 1/93/100 2 BRHt00 eata incerta insculpta umlaasiana olivania patula ocellata argus A argus B aterrima chamaeleon A = chamaeleon B HES histrio Q signata A signata B Amphinerita Argonerita 4100/100/100 peloronta 100/100/100 versicolor tata magdalenae Nerita s.s. 100/100/100 gx picea plicata A plicata B spenglenana A spengleriana B quadricolor grossa pec noes filosa 00/99/ oF IOSa A filosa C guamensis undulata A undulata B undulata C maxima undata (morph3) A pdate (norp oh g undata (mo, ~c undata 7p) A undata (morph1) = undata (incurva) A es una (maura) | ) com ‘a (nNovaeguineae earn Yagi on ata (ct novaeguineae 1198 helicinoides var. tristis Ritena ore po 10 Cymostyla Molecular phylogeny of Nerita species previously reconstructed from a combined Bayesian analysis of mitochondrial e estimated with Parsimony (MP) and Maximum Likelihood (ML) bootstrap proportions (BS = 70%), and Bayesian (MB) posterior probabilities (PP = 95%). Revised subgeneric groupings designated on far right. Asterisks indicate species that remain unplaced owing to their unstable phylogenetic positions. to illustrate the Nerita species-level phylogeny (Fig- ure 1). The phylogeny comprises each of the 11 recognized subgenera and 52 species (Table 1). NERITA SYSTEMATICS While the systematic relationships presented in Fig- ure | do not depart radically from previously proposed classifications within Nerita, the phylogenetic results suggest a need for minor revisions. At the tips of the phylogeny, the taxonomic status of previously recog- nized morphospecies was assessed using both molecular and morphological criteria. Nerita articulata Gould, 1847, N. antillarum Gmelin, 1791, N. georgina Récluz, 1841, and N. ornata Sowerby, 1833, appear to represent morphological variants rather than species (see Table 3a, Frey and Vermeij, 2008), and here they have been considered as such (Table 1). In contrast, N. tristis, M. A. Frey, 2008 Table 1 Nerita species included in the phylogenetic study (for detailed sample locations, see Frey & Vermeij, 2008). Due to unavailable tissue samples, the following species are not represented in the phylogeny, but they have been included in the revised classification: Nerita ascensionis Gmelin, 1791 (and subspecies N. a. chlorostoma Lamarck, 1816); Nerita dombeyi Récluz, 1841; Nerita fragum Reeve, 1855; Nerita luteonigra Dekker, 2000; Nerita nigrita R6oding, 1798; Nerita oryzarum Récluz, 1841; Nerita semirugosa Récluz, 1841; and Nerita sp. Eichhorst & Neville, In prep. Species and Author Nerita adenensis Mienis, 1978 Nerita albicilla Linnaeus, 1758 Nerita antiquata Récluz, 1841 Nerita argus Récluz, 1841 Nerita aterrima Gmelin, 1791 Nerita atramentosa Reeve, 1855 Nerita balteata Reeve, 1855 (includes articulata form) Nerita chamaeleon Linnaeus, 1758 Nerita costata Gmelin, 1791 Nerita erythrostoma Eichhorst & Neville, 2004 (syn. N. australis) Nerita exuvia Linnaeus, 1758 Nerita filosa Reeve, 1855 Nerita fulgurans Gmelin, 1791 (includes antillarum/N. lindae form) Nerita funiculata Menke, 1851 Nerita grossa Linnaeus, 1758 Nerita guamensis Quoy & Gaimard, 1834 Nerita helicinoides Reeve, 1855 Nerita histrio Linnaeus, 1758 (syn. N. squamulata) Nerita incerta von dem Busch in Philippi, 1844 Nerita insculpta Récluz, 1841 (includes georgina form) Nerita japonica Dunker, 1859 Nerita lirellata Rehder, 1980 Nerita litterata Gmelin, 1791 Nerita longii Récluz, 1842 Nerita magdalenae Gmelin, 1791 Nerita maxima Gmelin, 1791 Nerita melanotragus E. A. Smith, 1884 Nerita morio Sowerby, 1833 Nerita ocellata Le Guillou, 1841 Nerita olivaria Le Guillou, 1841 Nerita orbignyana Récluz, 1841 Nerita patula Récluz, 1841 Nerita peloronta Linnaeus, 1758 Nerita picea Récluz, 1841 Nerita planospira Anton, 1838 Nerita plicata Linnaeus, 1758 Nerita polita Linnaeus, 1758 Nerita quadricolor Gmelin, 1791 Nerita sanguinolenta Menke, 1829 Nerita scabricosta Lamarck, 1822 (includes ornata form) (auctorum) Nerita senegalensis Gmelin, 1791 Nerita signata Lamarck, 1822 (syn. N. reticulata) Nerita spengleriana Récluz, 1844 (syn. N. aurantia & N. oleagina) Page 3 Table 1 Continued. Species and Author Nerita tessellata Gmelin, 1791 Nerita textilis Gmelin, 1791 Nerita tristis Pilsbry, 1901 (syn. N. helicinoides var. tristis) Nerita umlaasiana Krauss, 1848 Nerita undata Linnaeus, 1758 Nerita undata (incurva form) von Martens in Kobelt, 1889 Nerita undata (maura form) Récluz, 1842 Nerita undata (novaeguineae form) Lesson, 1831 Nerita undata (striata form) Burrow, 1815 Nerita undulata Gmelin, 1791 Nerita versicolor Gmelin, 1791 Nerita vitiensis Hombron & Jaquinot, 1854 Nerita yoldii Récluz, 1841 previously recognized as a variant of N. helicinoides, has been elevated as a distinct species in light of the large genetic distance between the two taxa (Vermeij and Frey, 2008). Comprehensive sampling also re- vealed genetically distinct, evolutionary significant units (designated as lineage A, B, or C in Figure 1) within several species, including the N. undata complex (for sampling locales, see Table 3b in Frey and Vermeij, 2008). Additional study is necessary to determine whether these evolutionary lineages reflect distinct species or strong population genetic structure. At higher taxonomic levels, several clades nested within the genus generally correspond to previously recognized subgenera, indicating that the molecular findings agree with most morphology-based classifica- tions. However, some of the species, as formerly classified, result in polyphyletic clades; and the phylogenetic positions of a few species appear unstable across analyses (indicated by asterisks in Figure 1; for further details, see Frey and Vermeij, 2008). In order for the majority of subgeneric groupings to reflect monophyletic clades, I propose the classification shown in Table 2. While the relationships among the outlined groups remain uncertain, given the lack of strong support at deeper nodes, the majority of subgenera are well supported and, with the exception of Argonerita and Ritena, appear to be monophyletic. DISCUSSION Based on the molecular phylogeny, the two monotypic subgenera Adenerita and Ilynerita, represented by Nerita adenensis and N. planospira, respectively, appear to form a clade that falls as the sister taxon to all remaining Nerita species. As suggested by their monotypic assignments, both species exhibit shell morphologies that are distinguishable from all other extant Nerita spp., including each other (Dekker, 2000: Page 4 Table 2 Revised classification scheme of Nerita subgenera and species members. Type species indicated in boldface. Taxa not included in the presented phylogeny (denoted by asterisks) have been assigned to subgenera according to shell and opercular morphologies. Subgenera and Species Nerita sensu stricto Linnaeus, Ilynerita yon Martens, 1887 1758 Nerita peloronta Nerita scabricosta Nerita versicolor Adenerita Dekker, 2000 Nerita adenensis Amphinerita von Martens, 1887 Nerita incerta Nerita insculpta Nerita umlaasiana Argonerita Frey and Vermeij, 2008 Nerita argus Nerita aterrima Nerita chamaeleon Nerita fragum * Nerita histrio Nerita ocellata Nerita oryzarum * Nerita signata Cymostyla von Martens, 1887 Nerita filosa Nerita grossa Nerita guamensis Nerita helicinoides Nerita luteonigra * Nerita maxima Nerita nigrita * Nerita quadricolor Nerita semirugosa Nerita spengleriana Nerita tristis Nerita undata Nerita undulata * Heminerita von Martens, 1887 Nerita japonica Nerita yoldii Nerita planospira Linnerita Vermeij, 1984 Nerita antiquata Nerita erythrostoma Nerita litterata Nerita orbignyana Nerita polita Nerita vitiensis Lisanerita Krijnen, 2002 Nerita atramentosa Nerita lirellata Nerita melanotragus Nerita morio Ritena Gray, 1858 Nerita ascensionis * N. a. chlorostoma * Nerita costata Nerita magdalenae Nerita picea Nerita plicata Theliostyla Mérch, 1852 Nerita albicilla Nerita exuvia Nerita fulgurans Nerita funiculata Nerita sanguinolenta Nerita senegalensis Nerita tessellata Nerita textilis Nerita sensu lato (unplaced) Nerita balteata Nerita dombeyi * Nerita longii Nerita olivaria Nerita patula Krijnen, 2002). Despite shell differences, N. adenensis and N. planospira both possess an operculum with an unusually smooth outer surface (Mienis, 1978). If this trait represents a synapomorphy, then the long branches separating the two species reflect a deep divergence between sister taxa. Indeed, J/ynerita fossils indicate that any split between Adenerita and Ilynerita would have taken place prior to the late Eocene The Velisers V oles Nom (Vermeij et al., 2009). Alternatively, this shared trait may represent an ancestral condition, suggesting that the apparent sister relationship arises from long-branch attraction. This latter hypothesis rejects a sister species relationship, but it does not discount that these taxa represent basal lineages within the WNerita clade. Regardless of the exact relationship, I have retained each species in the respective subgenus, based on their genetic distance and morphological differences. The subtropical-temperate species of Nerita form two distinct monophyletic groups. The subgenus Lisanerita includes Nerita lirellata and N. morio, as originally proposed; however, phylogenetic results confirm that the clade also contains N. atramentosa and N. melanotragus (Spencer et al., 2007; Frey and Vermeij, 2008). These phylogenies also reveal that each respective pair of species, although remarkably similar in their shell morphologies, does not represent a set of sister species. Moreover, the lack of reciprocal mono- phyly between sister species N. morio and N. melano- tragus suggests a relatively recent split (see Waters et al., 2005; Spencer et al., 2007). These latter findings indicate strong and perhaps rapid convergence in shell morphology. Together, these Lisanerita lineages main- tain a basal, albeit unsupported, position across all analyses, suggesting an early divergence in the history of Nerita; however, the possible influence of long- branch attraction, pulling the clade towards the base of the tree, cannot be discounted. The monotypic Heminerita and Melanerita, as represented by Nerita japonica and N. yoldii, respec- tively, form a second: subtropical-temperate clade. Based on branch lengths, these species appear to be more closely related than previously recognized. Upon closer examination, N. japonica and N. yoldii show several similarities in morphology, including overall shell shape and, perhaps more importantly, an unusually large pseudoapophysis, which is an uncom- mon opercular trait within Nerita (Krijnen, 2002). To reflect the molecular and morphological affinities, I propose the placement of both species in the senior subgenus Heminerita. Although Heminerita forms a clade with the subgenus Theliostyla (as defined below) under certain phylogenetic analyses (results not illus- trated), few morphological characters support such an alignment. Both clades, however, maintain relatively basal positions, suggesting early splits within Nerita. Theliostyla, as traditionally recognized, represents a polyphyletic clade. The type species, Nerita albicilla, forms a strongly supported monophyletic group with N. sanguinolenta, which together unites with sister taxa N. exuvia and N. textilis. In turn, the group consistently aligns across all phylogenetic analyses with a well- supported clade that comprises N. funiculata, N. tessellata, and the closely related sister species WN. fulgurans and WN. senegalensis. Although the node M. A. Frey, 2008 uniting these two clades lacks strong support, these taxa possess similar shell and opercular features (Vermeij et al., 2009) and, with few exceptions, conform to previous descriptions of the subgenus (Krijnen, 2002). Accordingly, these species have been retained in Theliostyla. In contrast, several other species previously assigned to Theliostyla appear to form a distinct, more distantly related clade. Nerita chamaeleon, N. histrio, and N. signata form a well-supported clade that appears to be a sister group to N. aterrima, a species formerly placed in the subgenus Cymostyla. Together, this group consistently aligns with a monophyletic clade that comprises sister species N. argus and N. ocellata. To distinguish these species from Theliostyla, a new subgenus, Argonerita, was recently erected (see Appen- dix B, Frey and Vermeij, 2008). Based on the combined phylogeny, both N. patula and N. olivaria (see below) also align with this group; however, the respective nodes receive only limited support, and individual gene trees reveal alternative topologies (Frey and Vermeij, 2008). Consequently, I have designated these species as “unplaced” within Nerita sensu lato to reflect their unstable phylogenetic positions across analyses. While such assignments result in a nonmonophyletic Argo- nerita, the proposed classification scheme allows for future taxonomic changes to taxa that currently show uncertain phylogenetic placement. Two clades that are composed almost entirely of smooth-shelled species exist within Nerita. Amphinerita represents a strongly supported monophyletic clade that includes N. incerta, N. umlaasiana, and N. insculpta. Only N. insculpta departs from the smooth- shelled morphology, instead possessing narrow spiral ribs. Despite this difference, both N. umlaasiana and N. insculpta share several morphological features, includ- ing a distinctive crescent-shaped line that traverses the outer surface of the operculum (Krijnen, 2002). All three species are characterized by an exceptionally wide and glossy parietal callus. In addition to a completely smooth shell and a wide callus, N. olivaria also possesses a deep crescent-shaped opercular groove, leading many authors to classify this species as Amphinerita. Yet, surprisingly, the present molecular findings suggest otherwise. Conversely, N. balteata aligns with Amphinerita, but it shows few morpholog- ical similarities, apart from a superficial resemblance in shell sculpture with N. insculpta. Like N. olivaria, | have assigned N. balteata to Nerita sensu lato, based on its unstable phylogenetic position, and the striking incon- gruence between the molecular and the morphological evidence. Linnerita, the other smooth-shelled subgenus, repre- sents a well-supported monophyletic clade of six distinct species. The molecular findings correspond neatly to morphological and ecological traits, including Page 5 a characteristic smooth shell, a diagnostic operculum, and a strict association with sandy intertidal habitats (Vermeij, 1984). Within Linnerita, the type species N. polita forms a clade with N. orbignyana, while a sister clade comprises N. antiquata, N. vitiensis, N. litterata, and N. erythrostoma. The latter clade is also united by the presence of inconspicuous, fine, spiral cords (Eichhorst and Neville, 2004), which may reflect a synapomorphy for the group. Although the combined phylogeny reveals an alliance between Amphinerita, Linnerita, and the newly erected Argonerita, the relationships among these taxa remain obscured by unstable topologies and weak nodal support. As an aside, several other Nerita species also lack strong spiral sculpture (e.g., N. adenensis, N. senegalensis, N. maxima, N. morio, and N. picea), suggesting that a reduction in shell sculpture may have evolved repeat- edly in the genus. Species characterized by a high spire emerge as a clade (albeit with weak support), suggesting that a strong apex serves as a synapomorphy that unites Nerita sensu stricto, Ritena, and Cymostyla. The subgenus Nerita sensu stricto is represented by the type species N. peloronta. Although distinct shell and opercular features set this species apart from all other members of Nerita (Vermeij, 1984; Krijnen, 2002), NV. peloronta forms a _ well-supported clade with N. scabricosta and N. versicolor in each phylogenetic analysis. The long branches separating N. peloronta from its apparent sister species suggest a distant relationship. Indeed, a deep split, providing sufficient time for divergence, could explain differences in morphology. While the presented tree reveals UN. versicolor as the sister taxon, a separate gene tree shows N. scabricosta as the species most closely related to N. peloronta (Frey and Vermeij, 2008). Regardless of the sister-species relationship, both N. scabricosta and N. versicolor have been reassigned to Nerita sensu stricto to reflect the monophyly of the clade (Vermeij et al., 2009). With the transfer of these two species to Nerita sensu stricto, the subgenus Ritena appears to be a para- phyletic grade containing two groups. Nerita costata appears as the sister taxon to N. magdalenae; however, this relationship is neither stable nor supported across phylogenetic analyses (Frey and Vermeij, 2008). While these species resemble one another (Krijnen, 2002), the long branches separating the taxa indicate a distant split. Shell and opercular morphologies suggest that species of the unsampled N. ascensionis complex may serve as the true sister taxa to N. magdalenae. The other group represented by Ritena comprises the type species N. plicata and N. picea, a species previously assigned to Theliostyla. The molecular results consistently define these species as strongly supported sister taxa; however, apart from a high spire, these species show few Page 6 morphological similarities. In spite of the morpholog- ical differences, I have tentatively reassigned N. picea to Ritena until additional evidence suggests otherwise. While both groups contain at least one species characterized by a high spire, strongly sculptured spiral ribbing, a columella with transverse wrinkles, and a thin, concave operculum with fine granules (Dekker, 2000), Ritena possesses no obvious synapomorphies that set the clade apart from others, making the subgenus especially difficult to define (Krijnen, 2002). Despite paraphyly, I have retained both groups within Ritena until further investigation can verify clade membership. In contrast, the subgenus Cymostyla circumscribes two well-supported monophyletic clades. Within the first clade, N. quadricolor serves as sister species to a paraphyletic N. spengleriana. Together, these two species fall as sister taxa to a group that includes N. grossa, N. helicinoides, a paraphyletic N. filosa, and sister species N. guamensis. The second clade comprises N. undulata, N. maxima, representatives of the WN. undata complex, and N. tristis. Yet, based on shell morphology, both N. quadricolor and N. spengleriana show a striking similarity to members of the N. undata complex (Krijnen et al., 2006); and, as their previous nomenclature suggests, N. helicinoides differs only slightly from N. tristis (Vermeij and Frey, 2008). This discordance between the molecular phylogenetic results and the morphological characters suggests convergence in at least one of these groups. Overall, members of Cymostyla possess similar morphological features, including a high spire, distinct spiral ribbing, strong columellar teeth, and a coarsely granulated operculum. Based on these characters, previous classifications have also assigned N. longii to Cymostyla. In the reported phylogeny, N. Jongii aligns with Linnerita; however, the phylogenetic position of this species appears neither stable nor strongly supported across analyses. To reflect this uncertainty, I have transferred N. longii to Nerita sensu lato. The molecular phylogeny presented here substantial- ly clarifies our systematic understanding of WNerita. Additional molecular studies, including slower evolving markers, coupled with detailed anatomical and fossil studies, will be necessary to resolve the remaining phylogenetic uncertainties, particularly among basal nodes. Such advances are expected to reveal not only species and subgeneric relationships but also to improve knowledge of the evolutionary history of this prominent marine clade. Acknowledgements. A special thanks to the many who assisted with logistics and collection of specimens included in the phylogeny. H. Dekker, Y. Kano, C. Meyer, G. Paulay, D. Reid, and M. Schilthuizen provided additional tissue samples; D. Haasl, J. Harasewych, D. Reid, and K. Way granted access to museum specimens. I thank T. Eichhorst and G. Vermeij The Veliger, Vol. 51, No. 1 for insightful discussions, and T. Eichhorst, R. Grosberg, G. Vermeij, and P. Wainwright for thoughtful comments on the manuscript. This study was funded by research grants awarded to MAF from the Center for Biosystematics (UC Davis), Center for Population Biology (UC Davis), Conchol- ogists of America, Hawaiian Malacological Society, Lerner- Gray Fund for Marine Research (American Museum of Natural History), Polynesia Education and Research Labora- tories (UC Berkeley), Society of Systematic Biologists, and Western Society of Malacologists. Additional support was provided by grants from the National Science Foundation and the Mellon Foundation, awarded to R. Grosberg (IBN- 0416713 and OCE-06-23699) and G. Vermeij (EAR 97-06749). REFERENCES BERTNESS, M. D. & C. CUNNINGHAM. 1981. Crab shell- crushing predation and gastropod architectural defense. Journal of Experimental Marine Biology and Ecology 50: 213-230. CRANDALL, E. D., M. A. FREY, R. K. GROSBERG & P. H. BARBER. 2008. Contrasting demographic history and phylogeographical patterns in two Indo-Pacific gastro- pods. Molecular Ecology 17:611—626. DEKKER, H. 2000. The Neritidae (Gastropoda) from the circumarabian seas, with description of two new species, a new subgenus and a new genus. Vita Marina 47:29—64. EICHHORST, T. E. & B. D. NEVILLE. 2004. A review of the subgenus Linnerita Vermeij, 1984, (Gastropoda: Neriti- dae) with the designation of a new species and a redescription of Nerita (Linnerita) vitiensis Hombron & Jaquinot, 1854, from American Samoa and Fii. La Conchiglia 36:43-58. FRETTER, V. 1965. Functional studies of the anatomy of some neritid prosobranchs. Proceedings of the Zoological Society of London 147:46—74. Frey, M. A. & G. J. VERMEIJ. 2008. Molecular phylogenies and historical biogeography of a circumtropical group of gastropods (Genus: WNerita): implications for regional diversity patterns in the marine tropics. Molecular Phylogenetics and Evolution 48:1067—1086. GARRITY, S. D. 1984. Some adaptations of gastropods to physical stress on a tropical rocky shore. Ecology 65:559— 574. GRAY, J. E. 1858. Observations on the genus Nerita and its operculum. Proceedings of the Zoological Society of London 26:92-94. HOLTHUIS, B. 1995. Evolution between marine and freshwater habitats: A case study of the gastropod suborder Neritopsina. Zoology Department. University of Wash- ington, 249 pp. HurtTApbo, L. A., M. FREY, P. GAUBE, E. PFEILER & T. A. MaArKow. 2007. Geographical subdivision, demographic history and gene flow in two sympatric species of intertidal snails, Nerita scabricosta and Nerita funiculata, from the tropical eastern Pacific. Marine Biology 151: 1863-1873. Komatsu, S. 1986. Taxonomic revision of the Neritid gastropods. Special Publication of the Mukaishima Marine Biological Station, Pp. 1-69. KRIJNEN, C. 2002. The subgenera of the genus Nerita Linnaeus, 1758. Gloria Maris 41:38-69. KRIJNEN, C., F. CELEN, A. DELSAERDT, R. DUCHAMPs, L. NOTELTEIRS, N. REVIS, N. SEVERIJNS & M. VERHAEGHE. M. A. Frey, 2008 1995. Genus Nerita: alphabetical review, Part 1. Gloria Maris 34:65-84. KRIJNEN, C., F. CELEN, A. DELSAERDT, R. DUCHAMPS, L. NOTELTEIRS, N. REVIS, N. SEVERIJNS, M. VERHAEGHE & R. VINK. 1997. Genus Nerita: alphabetical review, Part 2. Gloria Maris 36:1—14. KRIJNEN, C., A. DELSAERDT & N. SEVERIJNS. 1999. Genus Nerita: alphabetical review, Part 3. Gloria Maris 38: 1-15. KRIJNEN, C., A. DELSAERDT, N. SEVERIJNS & M. VER- HAEGHE. 2001. Genus Nerita: alphabetical review, Part 4. Gloria Maris 40:27—52. KRIJNEN, C., A. DELSAERDT, N. SEVERIJNS & M. VER- HAEGHE. 2005. Genus Nerita: alphabetical review, Part 5. Gloria Maris 44:69-86. KRIJNEN, C., A. DELSAERDT, N. SEVERIJNS, M. VERHAEGHE & R. VINK. 2006. The problematic identity of Nerita undata Linne, 1758, with designation of a neotype (Gastropoda: Neritidae). Gloria Maris 45:66—90. LINNAEUS, C. 1758. Systema Naturae. Stockholm. MARTENS, E. V. 1887-1889. Die Gattungen Nerita und Neritopsis. In: W. Kobelt (ed.), Systematisches Conchy- lien-Cabinet von Martini und Chemnitz, Nurnberg, pp. 1— 64 (1887), 1865-1104 (1888), 1105-1147 (1889). MIENIS, H. K. 1978. Notes of recent and fossil Neritidae, 8. Nerita adenensis, a new species from the Arabian Peninsula. Argamon: Israel Journal of Malacology 6:30— 36. Morcu, O. A. L. 1852. Catalogus conchyliorum quae reliquit Page 7 D. Alphonso d’Aguirra & Gadea Comes de Yoldi, Hafnia: Copenhagen. SPENCER, H. G., J. M. WATERS & T. E. EICHHORST. 2007. Taxonomy and nomenclature of black nerites (Gastrop- oda: Neritimorpha: Nerita) from the South Pacific. Invertebrate Systematics 21:229-237. UNDERWOOD, A. J. 1984. Vertical and seasonal patterns in competition for microalgae between intertidal gastropods. Oecologia 64:211—222. UNDERWOOD, A. J. & M. G. CHAPMAN. 1996. Scales of spatial patterns of distribution of intertidal invertebrates. Oecologia 107:212—224. VERMEI, G. J. 1973. Morphological patterns in high- intertidal gastropods: adaptive strategies and their limita- tions. Marine Biology 20:319-346. VERMEI, G. J. 1984. The systematic position of the neritid prosobranch gastropod WNerita polita and related species. Proceedings of the Biological Society of Washington 97: 688-692. VERMEI, G. J. & M. A. FREY. 2008. Almost terrestrial: small supratidal species of Nerita (Gastropoda) in the western Pacific. Basteria 72:253-261. VERMEI, G. J.. M. A. FREY & B. M. LANDAU. 2009. The gastropod genus Nerita in the Neogene of tropical America. Cainozoic Research 6:61—70. WATERS, J. M., T. M. KING, P. M. O’LOUGHLIN & H. G. SPENCER. 2005. Phylogeographical disjunction in abun- dant high-dispersal littoral gastropods. Molecular Ecolo- gy 14:2789-2802. THE VELIGER IDs © CMS, Inc., 2008 The Veliger 51(1):8—14 (March 31, 2010) Bostrycapulus heteropoma n. sp. and Bostrycapulus tegulicius (Gastropoda: Calyptraeidae) from Western Africa RACHEL COLLIN* Smithsonian Tropical Research Institute, Apartado Postal 0843-03092, Balboa, Ancon, Republic of Panama AND EMILIO ROLAN Museo de Historia Natural, Campus Universitario Sur, 17582 Santiago de Compostela, Spain Abstract. Spiny slipper shells in the genus Bostrycapulus range worldwide in tropical and temperate oceans. Owing to the scarcity of samples that retain the defining characteristics of the genus, the species from tropical Africa and the Indo-Pacific are poorly known. Here we present data showing that samples of Bostrycapulus from the Cape Verde Islands and Senegal are distinct from each other and distinct from other known Bostrycapulus species. These two species can be distinguished from each other by the unique caplike protoconch found on the shells from Senegal and the coiled globose protoconchs typical of direct-developing species on the shells from the Cape Verde Islands. Genetically, samples from Senegal and the Cape Verde Islands are distinct. DNA sequences from Senegal are very similar to those of Bostrycapulus odites, which occurs in the South Atlantic (South Africa, Argentina, and Brazil) while those from Cape Verde are closest to Bostrycapulus aculeatus from Florida. The name Bostrycapulus tegulicius is available, and the single existing protoconch on the types in the Muséum National d’Histoire Naturelle, Paris, appears to match those from the Cape Verde Islands. Subtle variation in protoconch size and shape throughout the Cape Verde Islands suggests that there may be more than a single species in the archipelago. Unfortunately, too little material is available to rule out intraspecific variation or to support the description of additional new species from the Cape Verde Islands. Here, we augment the original description of B. tegulicius and describe the unique new species Bostrycapulus heteropoma n. sp. from Senegal. INTRODUCTION Cape Verde Islands is consistent with the description and type of B. tegulicius (Rochebrune, 1883). Observations of protoconch morphology, using both light microscopy and scanning electron microscopy (SEM), show distinct differences between the two species (Figures | and 2). Shells from Senegal have a cap-shaped protoconch that is attached at right angles to the teleoconch (Figure 1), a morphology that is unknown in any other calyptraeid, and which suggests an unusual mode of development. Animals from the Recent phylogenetic analyses of calyptraeid gastropods and a review of the genus (Collin, 2003a, b, 2005) have shown that there are at least eight species of Bostrycap- ulus that were previously synonymized under the name Crepidula aculeata or Bostrycapulus aculeatus (Gmelin, 1791; Hoagland, 1977, 1983). These species are difficult or impossible to distinguish with any confidence on the basis of adult shell characters or adult anatomy (Simone, 2002). However, they can generally be Cape Verde Islands have the coiled, globose proto- distinguished usin protoconch characters, develop- conchs typical of other direct developing Bostrycapulus mental observations, and DNA sequence data (Collin, (Collin, 2005). Examination of the one type of 2005). Samples that were recently collected from the Bostrycapulus tegulicius that retains a protoconch Cape Verde Islands and Senegal show that animals shows it to be similar to the material we obtained from these locations are different from each other and from Sao Vicente in the Cape Verde Islands (Figure 1). that the species from Senegal is distinct from previously Subtle variation in protoconch size and shape, but not described Bostrycapulus species. The species from the overall form, throughout the Cape Verde Islands suggests that there may be more than a single species in the archipelago (Figure 1). Alternatively, such *Corresponding author: Rachel Collin: collinr@si.edu; variation could be the result of Vvatiation ne TUES eee STRI, MRC 0580-08, Unit 9100 Box 0948, DPO AA34002, allocation or intraspecific variation in egg size. USA. Unfortunately, the mode of development of these R. Collin & E. Rolan, 2008 Figure 1. Bostrycapulus tegulicius. A. The lectotype of B. tegulicius that retains the protoconch. B. and C. SEMs of large globose protoconchs of B. tegulicius from Sal Island, Cape Verde. The arrow in B indicates the protoconch-teleoconch boundary and the arrow in C indicates the abrupt end of the radial sculpture. D. The tightly coiled protoconch of the lectotype of B. tegulicius. E. Protoconch of recently collected material from Sao Vicente. F. Globose protoconch from a pale shell from Sao Vicente. G. Globose protoconch from a dark shell from Sal. Scale bars: A = 5mm; B = 500 um; C = 450 um; D = 700 um; E = 750; F = 450 um; G = 450 um. species remains unknown, and too little material is available at this time to rule out intraspecific variation or to give strong support for the description of additional new species from the Cape Verde Islands. Cytochrome oxidase subunit I (COI) and 16S DNA sequence data obtained and analyzed following Collin (2000, 2001, 2005) show that samples from Senegal are not genetically similar to those from the Cape Verde Islands. Despite being geographically proximate, they fall into completely different parts of the phylogeny of the genus. The single live-collected sample that was available to us from Sal Island in the Cape Verde Islands is genetically distinct from other Bostrycapulus species (Figures 3 and 4). There is a 6—7% Kimura 2- Figure 2. Protoconchs of B. heteropoma from Senegal showing the distinctive protoconch morphology and the basis for the specific name heteropoma. B and E arrows point to protoconch. D and F arrows point to the ridge at the protoconch-teleoconch boundary. Scale bars: A = 500 um; B = 500; um C = D = 500 um; E = 500 um; F = 200 um. parameter divergence in COI sequences between this sample from the Cape Verde Islands and its sister species, B. aculeatus (Figure 3). A COI divergence of 6— 7% is similar to, or greater than, divergences between other distinct, well-recognized calyptraeid species (e.g., Collin, 2000, 2001, 2003b). The 16S sequences recover the same sister-species relationship but show lower levels of divergence (Figure 4), as is typical of this gene region. Bostrycapulus aculeatus has direct development with large, yolky eggs and a protoconch similar to those observed on shells from the Cape Verde Islands. Analysis of the COI and 16S sequences from animals collected in Senegal placed this species as sister to Bostrycapulus odites Collin 2005 from the South Atlantic, with only a 1.5—2.5% Kimura 2-parameter divergence in COI. This is a small COI divergence for sister species, but some other morphologically distinct calyptraeid species with similar or smaller interspecific distances have been reported (Collin, 2003b; Collin et al., 2007). Bostrycapulus odites has direct development Page 10 SS B. heteropoma 100 re B. odites 63 B. urraca 100 94 B. calyptraeformis 100 100 98 100 B. latebrus B. pritzkeri B. gravispinosus 100 B. aculeatus 100 B. tegulicius — 0.005 substitutions/site Figure 3. Neighbor-joining tree of Bostrycapulus based on 611 base pairs of mt COI DNA sequence data. All data are from Collin (2005) except for three individuals of B. heteropoma. Numbers above the branches show bootstrap support for the major nodes. More details can be found in Collin (2005). from small eggs, where the embryos consume other eggs and embryos, and it has a coiled protoconch that is clearly different from those from Senegal. The unique protoconch on shells from Senegal (Figure 2), which is unlike the protoconch of any other Bostrycapulus species (Collin, 2005) and is, in fact, of a type unknown for any other calyptraeid, leaves no doubt that this is a distinct species. The Veliger, Vol. 51, No. 1 B. heteropoma B. odites B. urraca B. calyptraeformis B. latebrus B. pritzkeri B. gravispinosus B. aculeatus B. tegulicius == 0.005 substitutions/site 16S Figure 4. Neighbor-joining tree of Bostrycapulus based on 481 base pairs of mt 16S DNA sequence data. All data are from Collin (2005) except for three individuals of B. heteropoma. Numbers above the branches show bootstrap support for the major nodes. More details can be found in Collin (2005). SYSTEMATICS Genus Bostrycapulus Olsson & Harbison, 1953 Type Species Bostrycapulus aculeatus (Gmelin) by original designation Bostrycapulus tegulicius (Rochebrune, 1883) (Figure 1) Synonymy: Crypta tegulicia Rochebrune 1883:4. Crepidula aculeata (Gmelin, 1791), Lamy 1911:318 (in part); Hoagland 1977:364 (in part); Hoagland, 1983:7 (in part). Crepidula cf. tegulicia Collin, 2003a: 541-593. Collin, 2003b:618-640. Bostrycapulus cf. tegulicius Collin, 2005. Types: Two syntypes are in the Muséum National d’Histoire Naturelle, Paris (MNHN) (Hoagland, 1983). R. Collin & E. Rolan, 2008 Both figured in Hoagland (1983, fig. 9). The shell figured as 9a in Hoagland (1983) retains a protoconch (Fig- ure 1). We hereby designate this shell as the lectotype of Bostrycapulus tegulicius. The shell length is 17.6 mm. Original description: “Testa subovata, crassiuscula, irregulari, oblique curvata, extus albida, concentrice striata, et squamis minutis teguliformibus, subdistanti- bus orniata; intus nitide castaneo violacea; lamella opalina, ad medio et ad latus subemarginata. Long 0.019, Lat 0.014.” Type locality and distribution: The original description cites ““‘Dakar, Joalles, Pointe de Cap Vert’ Senegal as the locality. However, all of the Bostrycapulus shells collected by one of us (ER) in Senegal retain protoconchs that demonstrate them to be distinct from B. tegulicius. Material collected from the Cape Verde Island on Sao Vicente Island matches the protoconch morphology of the type of B. tegulicius (Figure 1). Therefore, we believe that the locality cited by Rochebrune (1883) is in error. This is likely, since type localities from the nineteenth century are often approximate, and Rochebrune examined material collected by a variety of people and is unlikely to have examined the protoconchs in detail. Habitat: Most of the material examined was collected by dredging at 30-40 m, and living material was most often found on dead bivalve shells on soft bottoms. One animal was collected from the shell of a living Conus sp. from 2 m depth. Material examined: Cream morphotypes with gracile protoconchs (B. tegulicius s. s.): 2 juveniles, 3 females, Sao Vicente, Cape Verde (MNHN). 2 specimens, Porto Mindelo, Sao Vicente, Cape Verde, 15m (MHNS : Museo de Historia Natural of the University of Santiago de Compostela). 1 shell, Matiota, Sao Vicente, Cape Verde (MHNS). Material examined: Brown morphotypes with globose protoconchs: 5 juveniles, Santa Maria, Sal, 30 m (MHNS). 1 shell, Algodoeiro, Sal, Cape Verde (MHNS). 3 shells, Palmeira, Sal, Cape Verde, 30 m (MHNS). 1 shell, Pau Seco, Maio, Cape Verde, 30 m (MHNS). 50 shells, 17 juveniles, Sal, Cape Verde (MNHN, coll. Marche-Marchad). 1 shell, Boavista, Cape Verde (coll. Michele) (the shell _ represented in Ardovini & Cossignani, 2004:100). 1 juvenile, Sal, Cape Verde, FMNH282359 Diagnosis: Bostrycapulus tegulicius can be distinguished from all other known species of Bostrycapulus by a combination of protoconch morphology, DNA se- quence data, and biogeography. The protoconch has Page 11 a single 850-1000 um whorl, available sequence data for the DNA barcoding gene COI, GenBank #tAY061776, is distinct from other species, and this species is known to occur only in Cape Verde. Description: Examination of dead material from several localities throughout the Cape Verde Islands shows that there is subtle variation both in shell color and in protoconch size (cream and brown morphotypes above) and that this variation may represent intraspe- cific or interspecific variation (see below). The follow- ing description is based on samples (cream morpho- types) that match the protoconch of the type of B. tegulicius and that are generally typical of samples from Sao Vicente. Bostrycapulus tegulicius sensu stricto (cream mor- photype) has adult morphology typical of all Bostry- capulus species. The spiny shells, up to 25 mm in length, are convex and retain distinct coiling. There is a lunate muscle scar anterior to the septum on the animal’s right side, and the edge of the septum is sinuous (see Collin, 2005, for detailed description). The shells of B. tegulicius are generally pale with sparse brown markings and can be distinguished from the shells of other species of Bostrycapulus by protoconch morphology. The protoconch is large and globose, and it constitutes about one whorl, with a length from the posterior of the shell to the protoconch—teleoconch boundary of about 850-1000 um (Figure 1E). The protoconch-teleochonch boundary is not clearly de- marcated. The samples available to us are all uniformly smooth, and there is no indication of the fine, granular sculpture that is present on the shells of several other Bostrycapulus species. Many of the shells we examined were slightly eroded. Because the granular sculpture on the shells of other species erodes easily (Collin, 2005), it is possible that newly hatched individuals of this species have granular sculpture. No embryos have been observed in live-collected material, and therefore the details of embryonic development and egg-size mea- surements are not available. Neither DNA sequence data nor appropriate tissue for molecular work for samples that unambiguously match the type are available at this time. Individuals of the brown morphotype of Bostrycap- ulus tegulicius have brown shells with cream markings that are typical of samples from Sal Island and Boavista Island. They have larger, more globose protoconchs (Figure 1B) than the pale morphs from Sao Vicente. The protoconch is large, globose, and constitutes about half a whorl with a length from the posterior of the protoconch to the protoconch— teleoconch boundary of about 800-900 um (Figure 1). The protoconch-teleoconch boundary is_ clearly marked by the abrupt initiation of spiral cords (Figure 1). The spiral cords give way to an underlying Page 12 smooth sculpture with occasional plicate spines after an additional 800-900 um. There is no indication of fine granular sculpture. The protoconch morphology and the DNA sequenc- es described in Collin (2005) and attributed to B. cf. tegulicius correspond to the brown morphotype of B. tegulicius. In general, protoconchs from darker shells are slightly larger and less coiled than those from pale shells (Figure 1). However, shells with similar globose protoconch morphologies have also been observed on pale shells from Sao Vicente (e.g., Figure 1F), demon- strating that the general association between shell color, protoconch morphology, and locality is not rigid. Development: unknown. The protoconch morphology is consistent with direct development from large yolky eggs. Sequences in GenBank: COI AY061776, 16S AY061775 for FMNH282359 Sal Island, Cape Verde. Remarks: The taxonomic status of the two Bostrycap- ulus morphotypes present in Cape Verde is not clear. It is clear that the pale morphotype with the more coiled protoconch matches the type of B. tegulicius. However, whether differences in protoconch morphology and/or shell color indicate the presence of one or several additional species is unclear. In general, the proto- conchs from darker shells are larger and less coiled than those from pale shells. However, all combinations of color and protoconch morphology do occur. Some of these differences may simply be the result of geographic variation within a species, as shell color and sculpture varies among populations of other Bostrycapulus species (Collin, 2005). Alternatively, it is possible that these differences indicate the presence of two very closely related species. The Cape Verde Islands are the site of very recent explosive radiations of other marine gastropods with direct development (e.g., Conus [Duda & Rolan, 2005; Cunha et al., 2005]) and therefore interspecific divergences may also be very recent. Because protoconch morphology usually varies little within a species, we anticipate that distinct specific status of animals with each protoconch morphology will be supported as more material becomes available for study. However, we prefer the conservative approach of waiting until more than one line of evidence is available to support the introduction of additional species names. Bostrycapulus heteropoma Collin & Rolan, sp. nov. (Figures 2, 5) Synonymy: Crepidula aculeata (Gmelin, 1791) Adam & Leloup, 1935:358 (in part). Hoagland 1977:364 (in part). The Veliger, Vol. 51, No. 1 Figure 5. The holotype of Bostrycapulus heteropoma MNHN 21222. A. Dorsal view of the shell. B. and D. Different views of the protoconch. C. Ventral view of the holotype showing the tissue that was removed for sequencing. Hoiotype: Gorée, Dakar, Senegal. Individual #282. GenBank #-DQ314567; DQ314570. Muséum National d’Histoire Naturelle, Paris. MNHN 21222. Figure 5. Shell length = 25.5mm. Paratypes: Gorée, Dakar, Senegal. Individual #284. GenBank #DQ314569..ANSP A21823. Shell length = 20.6 mm. Type locality: Gorée, Senegal. 14°24’N 19°30'W. Other Material Examined: One shell, Gouye Teni M’Both, Dakar, Senegal, 25 m (J. Pelorce collection, Paris). One shell, Cap Vert, Dakar, Senegal (MHNS). Three juveniles, Goute Teni M’Both, Dakar, Senegal, 28 m (J. Pelorce collection, Paris). One shell, Petit Corniche, Dakar, Senegal, 40 m (J. Pelorce collection, Paris). Three specimens, Cap Vert, Dakar, Senegal, 40 m (J. Pelorce collection, Paris). Eight juveniles, Gorée, Dakar, Senegal, 8—15 m (MHNS). One juvenile, N’Gor, Dakar, Senegal, 7 m (MHNS). One female, Gorée, Dakar, Senegal. Individual #283. GenBank #DQ314568. ANSP A21822; shell length = 25.5 mm; shell very eroded by epibionts. Distribution and Habitat: This species has been collected from Dakar, Senegal. The material examined here was collected attached to large rocks from subtidal sandy habitats. R. Collin & E. Rolan, 2008 Diagnosis: Bostrycapulus heteropoma can be distin- guished from all other known species of Bostrycapulus by its unique protoconch morphology, its distribution in Senegal, and the sequence of the COI gene. The cap- shaped protoconch is slightly curved and attaches to the teleoconch at a right angle along a prominent ridge. Description: The adult shell morphology and anatomy are typical of Bostrycapulus species (see Collin, 2005, and Simone, 2002, for detailed descriptions). External- ly, the shell is relatively flattened and somewhat coiled. The internal septum extends about half the length of the shell, and the anterior margin is indented medially and notched on the animal’s left. There is a distinct but small medial ridge or crease from the medial indenta- tion to the posterior shell margin near the apex. There is a small lunar muscle scar on the animal’s right side anterior to the septum. The apex is appressed, usually slightly above the posterior shell margin on the right and is not excavated. The external shell sculpture varies from widely spaced, large, scalelike plicate spines to tightly packed, pointed, granular bumps along fine spiral ribs. Shell color varies from overall cream with scattered brown markings to solid chocolate brown, sometimes with pale streaks and occasionally solid tan. Markings are sometimes speckled and often streaky. The protoconch is a large, elongate cap with pointed, asymmetrical apex and shows a slight curve but little sign of coiling (Figures 2 and 5). The 700 um proto- conch is attached at right angles to the teleoconch, and the boundary is marked by a distinct ridge (Figures 2 and 5). This morphology is so distinct that, without direct observations of development, it is not possible to verify that this ridge does indeed represent the protoconch-teleoconch boundary. We decide to call the ridge the protoconch—teleoconch boundary simply because there are no other clear demarcations on the shell that could be interpreted in this way, and because most calyptraeid shells include a clear protoconch— teleoconch boundary. It is possible that the ridge represents the boundary between protoconch I, laid down by the embryo prior to ingestion of nurse eggs or intracapsular albumin, and that the smooth area after the ridge is a later stage of a protoconch (protoconch II). However, if this is the case, there is no clear protoconch-teleoconch boundary. The protoconch is smooth, with some indication of growth lines prior to the ridge. The shell has more distinct growth ridges and some indistinct radial sculpture subsequent to the ridge. Some specimens retain very fine granular sculpture on the early teleoconch (Figure 2). The pigmentation of the living animals is similar to that of other Bostrycapulus species. The body is generally creamish in large animals and translucent in very small animals. In large animals, the foot is a darker yellowish and the head is often orange. The Page 13 tentacles and snout have creamish or yellowish pigment splotches, and the mantle is decorated with numerous small, irregular white or cream spots. The osphradium is bipectinate with a brown border. Unlike other Bostrycapulus species, the animals we examined did not have black pigment along the edge of the foot or on the head or neck. We did not detect any diagnostic differences in the internal anatomy of ethanol-pre- served samples. However, there were generally more distinct lobes on the capsule gland and the albumin gland than are usually observed in other Bostrycapulus species. Development: Unknown. Sequences in GenBank: COI: 3 sequences, GenBank # DQ314567-69; 16S: 1 sequence, GenBank # DQ314570. Etymology: The neuter Greek noun heteropoma means “strange cover” and refers to this species’ unusual protoconch morphology. Acknowledgments. We thank José Hernandez for field assistance in Dakar, where the new species was collected; P. Bouchet and Virginie Héros of the MNHN for the loan of the type of Crypta tegulicia and for access to collections under their care; M. Venegas for DNA sequencing; Jesus Méndez, of the CACTI, University of Vigo, for the SEM photos; and Jacques Pelorce and Michela Kuan for access to their collections. LITERATURE CITED ADAM, W. & E. LELoup. 1935. Les Crepidula de la Cote Occidentale de l’Afrique. Mémoires du Musée Royal d’Histoire Naturelle de Belgique, Ser 2., 3:347—367. ARDOVINI, R. & T. COSSIGNANI. 2004. West African Seashells. Mostra Mondiale Malacologia, 319 pp. COLLIN, R. 2000. Phylogeny of the Crepidula plana (Gastrop- oda: Calyptraeidae) cryptic species complex in North America. Canadian Journal of Zoology 78:1500—1514. COLLIN, R. 2001. Effects of mode of development on phylogoegraphy, and population structure of North Atlantic Crepidula (Gastropoda: Calyptraeidae). Molecu- lar Ecology 10:2249-2262. COLLIN, R. 2003a. The utility of morphological characters in gastropod phylogenetics: An example from the Calyp- traeidae. Biological Journal of the Linnean Society 78: 541-593. COLLIN, R. 2003b. Phylogenetic relationships among calyp- traeid gastropods and their implications for the bio- geography of speciation. Systematic Biology 52(5):618— 640. COLLIN, R. 2005. Development, phylogeny, and taxonomy of Bostrycapulus (Caenogastropoda: Calyptraeidae), an an- cient cryptic radiation. Zoological Journal of the Linnean Society 144:75-101. COLLIN, R., O. R. CHAPARRO, F. WINKLER & D. VELIZ. 2007. Molecular phylogenetic and embryological evidence that feeding larvae have been reacquired in a marine gastropod. Biological Bulletin 212:83—92. CUNHA, R. L., R. CASTILHO, L. RUBER & R. ZARDOYA. 2005. Page 14 Patterns of cladogenesis in the venomous marine gastro- pod genus Conus from the Cape Verde Islands. Systematic Biology 54(4):634-650. DubDaA, T. F. & E. ROLAN. 2005. Explosive radiation of Cape Verde Conus, a marine species flock. Molecular Ecology 14(1):267-272. GMELIN, J. F. 1791. Systema Naturae, ed 13, 1(6):3021-4120. HOAGLAND, K. E. 1977. Systematic review of fossil and recent Crepidula and discussion of evolution of the Calyptraei- dae. Malacologia 16(2):353-420. HOAGLAND, K. E. 1983. Notes on type specimens of Crepidula (Prosobranchia: Calyptraeidae) in the Muséum National d’Histoire Naturelle, Paris. Proceedings of the Academy of Natural Sciences, Philadelphia 135:1-8. Lamy, E. 1911. Sur quelques Mollusques de Sénégambie. The Veliger, Vol. 51, No. 1 Bulletin de Muséum National d’ Histoire Naturelle (Paris) 17:316-319. Quoy, J. R. C. & J. P. GAIMARD. 1832-1833. Voyage de lAstrolabe. Zoologie, Mollusca 2:1—686. ROCHEBRUNE, A. T. de. 1883. Diagnosis de mollusques nouveaux propres a la Sénégambie. Bulletin de la Société Philomathique de Paris. Ser. 7:177—182. SIMONE, L. R. L. 2002. Comparative morphological study and phylogeny of representatives of the superfamilies Calyptraeoidea (including Hipponicoidea) (Mollusca, Caenogastropoda). Bioto Neotropica 2(2), http://www. biotaneotropica.org.br. WERNER, B. & K. G. GRELL. 1950. Die amerikanische Pantoffelschnecke Crepidula fornicata L.: Eine Anleitung zur Préparation. Verlag von Gustav Fischer: Jena. 24 pp. The Veliger 51(1):15—25 (March 31, 2010) _ THE VELIGER © © CMS, Inc., 2008 A New Species of Hypselodoris and a Redescription of Hypselodoris picta lajensis (Nudibranchia: Chromodorididae) from Brazil SIMONE DACOSTA Zoologische Staatssammlung Miinchen, Minchhausenstr. 21, 81247 Minchen, Germany (e-mail: simonedacosta@gmx.de) VINICIUS PADULA Departamento de Invertebrados, Museu Nacional / Universidade Federal do Rio de Janeiro, Quinta da Boa Vista s/n, Sao Cristovao, 20940-040, Rio de Janeiro, RJ, Brasil (e-mail: viniciuspadula@yahoo.com) MICHAEL SCHRODL Zoologische Staatssammlung Miinchen, Minchhausenstr. 21, 81247 Miinchen, Germany (e-mail: schroed1@zi.biologie.uni-muenchen.de) Abstract. The external and internal morphology of Hypselodoris juliae sp. nov. is described in detail. This beautiful new species differs from congeners by its pale notum with many orange or yellow lines and iridescent blue patches, a relatively short prostate, and the absence of denticles on the lateral radular teeth. Its known geographic distribution includes the southeastern Brazilian coast, and possibly the Caribbean Sea. Tropical western Atlantic species of Hypselodoris are discussed comparatively. A redescription of Hypselodoris picta lajensis, which is known only from the southeastern and southern coast of Brazil, is presented. For the first time, wide intraspecific variation in coloration, radula, and features of the reproductive system of this subspecies was observed. Because of this variation, we prefer to maintain the subspecies rank of H. picta lajensis until a comprehensive revision of all H. picta material, including detailed anatomical comparisons and also a molecular approach, can provide a better understanding of this group. INTRODUCTION Most of the knowledge of the Brazilian nudibranch fauna results from the studies of Ernst and Eveline Marcus between 1950 and 1980. From samples in the intertidal zone and from material obtained from research vessels, those authors described most of the Brazilian species (Ev. Marcus, 1977; Rios, 1994). Scuba diving now enables researchers to collect specimens in the still poorly studied Brazilian subtidal zone. Consequently, a number of species have recently been discovered and described from offshore islands and the continental coast (Troncoso et al., 1998; Garcia et al., 2002; Garcia & Troncoso, 2003, 2004; Pola et al., 2005; ‘Dominguez et al., 2006a, b). Our own investigations on aeolid nudibranchs resulted in four new records from the Brazilian coast (Padula & Absalao, 2005; Padula & Santos, 2006) and in the description of a new Brazilian subspecies of Flabellina engeli Marcus & Marcus, 1968 (DaCosta et al. 2007). To date, approximately 100 nudibranch species have been reported from Brazil. Three of them belong to the genus Aypselodoris: Aypselodoris marci Marcus, 1971; H. picta lajensis Troncoso et al., 1998, and H. sycilla (Bergh, 1890). This number is very small, since the genus currently comprises at least 69 species, most of which occur in tropical and subtropical seas (Wilson & Willan, 2007; Ortea & Bacallado, 2007). In the present study, we describe in detail the external and internal morphology of a new species of Hypselodoris collected at Cabo Frio, on the southeast- ern coast of Brazil. The new species is compared with congeners from Brazil and the Caribbean Sea. Hypse- lodoris picta lajensis, for which major anatomical data was basically limited to the type specimen, is compre- hensively redescribed. MATERIAL AND METHODS The specimens were collected manually in the intertidal zone and through scuba diving on different sites in the Page 16 Cabo Frio region, on the southeastern Brazilian coast, between December 2002 and December 2006. In the laboratory, the specimens were photographed and measured alive, relaxed with a 10% MgCl, solution, and preserved in 70% ethanol. With the aid of a binocular microscope, two specimens of Hypselodoris juliae sp. nov. and six specimens of H. picta lajensis were dissected, each through a dorsal incision. Internal features were examined and drawn using the camera lucida of the microscope. The buccal mass was removed and dissolved in a 10% sodium hydroxide solution to isolate the armed labial cuticle and the radula from the surrounding tissue. Then the hard parts were rinsed in water, dried, and mounted for examination with a scanning electron microscope (SEM). Abbreviations: MNRJ, Museu Nacional/Universidade Federal do Rio de Janeiro; ZSM, Zoologische Staats- sammlung Minchen. SYSTEMATICS Family CHROMODORIDIDAE Bergh, 1891 Genus Hypselodoris Stimpson, 1855 Type species: By monotypy, Goniodoris? obscura (Stimpson, 1855). Port Jackson, Australia. Hypselodoris juliae sp. nov. (Figures 1-4) ? Hypselodoris sp. 4 Rudman, (2001—2003). ? Hypselodoris sp. 1 Valdés et al., 2006: 165. Material examined: Holotype: MNRJ 10940, 57 mm alive, dissected, Praia das Conchas, Cabo Frio, state of Rio de Janeiro, Brazil, 22°52'15”S, 41°58'52”W, intertidal, 14 Novem- ber 2004, Jeg. V. Padula. Paratype: ZSM 20040149, 1 specimen, dissected, Ilha dos Papagaios, Cabo Frio, state of Rio de Janeiro, Brazil, 22°53'49"S, 41°59'2”W, 8 m depth, 28 Decem- ber 2002, leg. S. DaCosta. Additional material: A color photo of a living specimen from Guarapari, state of Espirito Santo, Brazil. Etymology: This species is dedicated to Julia Carvalho Schroédl, the daughter of Michael Schrédl. Geographic distribution: Southeastern coast of Brazil: state of Rio de Janeiro (Cabo Frio region) and the state of Espirito Santo (Guarapari). Note: specimens from the Caribbean region illustrated by Rudman (2001-— The Veliger, Vol. 51, No. 1 2003) as Hypselodoris sp. 4 and by Valdés et al. (2006) as Hypselodoris sp. 1 appear to be conspecific. Anatomical studies on specimens from these localities will be necessary to confirm this distribution. External morphology: Living holotype 57 mm long. Body elongate, slightly higher than wide. Background color pale blue to pale green with small iridescent blue patches, and covered by numerous parallel longitudinal yellow or orange lines (Figure 1A). Lateral body wall and posterior dorsal region of foot with similar pattern, becoming pale blue with orange spots near the foot sole (Figure 1C). Genital opening surrounded by an orange ring. Foot sole pale blue (Figure 1B). Holotype (MNRJ 10940) with 21, and paratype (ZSM 20040149) with 13, continuous dorsal lines plus some incomplete lines running from anterior to posterior notum margin, interrupted by rhinophores and gill circle. Orange line surrounding the base of each rhinophore. Region above the eyes without pigmenta- tion. Undulating mantle border; margin orange on dorsal and ventral side. Submarginal grayish band with pale blue circular spots, except for dark middle notum area with two short transverse orange lines (Fig- ure 1A). Anterior and posterior notal spots encircled by orange lines. Row of pale blue spots on the submarginal band corresponding to the position of mantle dermal formations (MDFs). No MDEFs in dark middle area. Bilabiated anterior foot margin, upper lip orange, not notched. Oral tentacles short, digitiform, very near the mouth. Rhinophores retractile into the moderately elevated sheaths. Rhinophores perfoliate with 21 lamellae (holotype: MNRJ 10940) and 19 lamellae (paratype: ZSM 20040149); dark green with a dark-blue distal portion in the holotype and uniformly dark blue in the paratype; rhinophores with clear tip and a posterior small white mark near the base. Low gill sheath; longitudinal orange lines ending at margin, i.e., without encircling gills. Ten unipinnate, bipinnate, or tripinnate yellowish gills (Figures 1A and 1C). Gill rachis black in the holotype and dark blue in the paratype, with orange margins. Rows of small whitish spheres, probably glands, shining through gill tissue. Digestive system: Oral tube wide and strong, increasing in volume after a slight constriction at its midlength (Figure 3A). Jaw rodlets stout to elongate, unicuspid. Bulbous pharynx with posteroventrally projecting radula sac. Radula measuring 4mm in length in holotype, with formula 85 xX 165.0.165. Radula of paratype measuring 2.6 mm in length, with formula 58 X 125.0.125. Rachis narrow, devoid of teeth. Lateral teeth hook-shaped, bicuspid; slender cusps, pointed, almost equally long or upper cusp slightly longer than lower cusp. Some initial lateral teeth of holotype having only vestigial upper cusps. Outer lateral teeth elongate with very short, blunt cusps (Figure 2). S. DaCosta et al., 2008 Page 17 Figure 1. Aypselodoris juliae sp. nov., living holotype (MNRJ 10940): A, dorsal view; B, ventral view; C, lateral view. Hypselodoris picta lajensis (MNRJ 11973), intraspecific variation: D, adult specimen with many dorsal lines, yellow mantle margin; E, adult specimen with three dorsal lines and yellow mantle margin. Tubular salivary glands, entering pharynx laterally to thin-walled, tubular esophagus. Stomach and spherical calcum completely embedded in the holohepatic digestive gland. Intestine leaving stomach posteriorly, then bending upward and forward, leaving digestive gland as a slender tube. Middle portion of intestine more voluminous, forming a loop directed backward to right side of body; narrow distal intestinal portion, running toward the anal papilla in the center of the gill circle (Figure 3A). Page 18 The Veliger, Vol. 51, No. 1 Figure 2. Hypselodoris juliae sp. nov., SEM micrographs of the holotype (MNRJ 10940): A, jaw rodlets; B, innermost radular teeth; C, midlateral teeth of the radula; D, outermost radular teeth. Circulatory system: Voluminous pericardium lying posteromedially in the body cavity above gonad and digestive gland. Two-chambered heart, longitudinally oriented; trapezoidal auricle, posterior to muscular, spherical ventricle. Aorta dividing into posterior aorta running to left side of viscera and anterior aorta leading to the head (Figure 3B). Whitish blood gland, consist- ing of two flat lobes; posterior lobe partly covering salivary glands and distal reproductive system, anterior lobe very elongate and reaching to anterior portion of body cavity. Syrinx leaving the pericardium ventrally to the atrium (Figure 3B). Nervous system: Comprising paired cerebropleural, pedal, buccal, gastroesophageal, and _ rhinophoral ganglia, which surround anterior esophagus (Fig- ure 4A and 4B). Cerebral and pleural ganglia com- pletely fused, i.e., no external separation detectable; single, short, thick commissure. Cerebropleural ganglia much larger than pedal ganglia. Cerebropleural—pedal connectives very short. Statocyst with several otoconia laterally attached between cerebropleural and pedal ganglia. Small rhinophoral ganglia anteriorly attached to cerebropleural ganglia. Single rhinophoral nerve leaving each rhinophoral ganglion and leading to the accessory ganglion within rhinophore stalk; some thin nerves innervating rhinophoral lamellae. Pigmented eyes, short optic nerve. Buccal ganglia situated between the radula sac and the esophagus. Very short buccal commissure. Small gastroesophageal ganglia (Fig- ure 4B). Reproductive system: Gonad covering digestive gland dorsally. Proximal gonoduct a long thin tube. Distal S. DaCosta et al., 2008 A Page 19 ey gen Figure 3. Aypselodoris juliae sp. nov., schematic anatomical drawings: A, digestive system; B, organization of major organ systems. Abbreviations: aa, anterior aorta; ao, anus opening; ap, posterior aorta; bgl, blood gland; ca, digestive caecum; dg, digestive gland; ey, eyes; gen, reproductive organs; in, intestine; mo, mouth; oe, esophagus; ot, oral tube; ph, pharynx; r, radula: rhs, rhinophoral sheath; sg, salivary gland; st, stomach; ve, ventricle. genital system triaulic. Hermaphroditic ampulla short and swollen. Short distal gonoduct. Thin oviduct, entering female gland mass after a short distance. Short prostate. Prostatic loops attached to bursa copulatrix. Distal vas deferens a winding tube, relatively short. Ejaculatory duct simple, without armature, slightly everted into the elongate penial sheath (Figure 4C). Male and female atria distally combined into common vestibule. Vagina a relatively wide tube. Allosperm receptacles vaginally arranged. Bursa large, spherical, thin-walled. Receptaculum seminis club-shaped, half the diameter of bursa, short-stalked. Thin uterine duct splitting off vagina slightly distal to, and opposite of, receptacle stalk; uterine duct entering female gland mass close to the oviduct. Female gland mass well ‘developed, consisting of several barely distinguishable glandular portions. Nidamental duct showing conspic- uous unilateral swelling; nidamental opening posterior to joint vestibule. Flat glandular lobe covering half of female gland mass; deferent duct hidden between nidamental opening and vestibule (Figure 4C). Hypselodoris picta lajensis Troncoso, Garcia, & Urgorri, 1998 (Figures 5—7) Hypselodoris picta lajensis Troncoso, Garcia, & Ur- gorri, 1998: 135 Hypselodoris lajensis Troncoso, Garcia, & Urgorri, 1998: Dominguez, Garcia & Troncoso (2006: 633) Material examined: ZSM 20040122, one specimen, 20 mm fixed, dissected, Ilha do Papagaio, Cabo Frio, state of Rio de Janeiro, Brazil, 22°53'49”S, 41°59'2’/W, 28 December 2002, 10 m depth, Jeg. S. Da Costa. ZSM 20040123, one specimen, 25 mm fixed, dissected, Ilha do Papagaio, Cabo Frio, state of Rio de Janeiro, Brazil, 22°53’49"S, 41°59’2”W, 28 December 2002, 6 m depth, Jeg. S. DaCosta. MNRJ 11972, two specimens, 35-50 mm alive, dissected, Ilha Comprida, Cabo Frio, state of Rio de Janeiro, Brazil, 22°52’15”S, 41°56'53’W, a The Veliger, Vol. 51, No. 1 Figure 4. Hypselodoris juliae sp. nov. A, B, central nervous system; C, reproductive system. Abbreviations: am, ampulla; be, bursa copulatrix; bg, buccal ganglia; epl, cerebropleural ganglia; de, ejaculatory duct; en, optic nerve; ey, eye; fgm, female gland mass; goe, gastroesophagial ganglia; hd, hermaphrodite duct; nd, nidamental duct; no, nidamental duct opening; ov, oviduct; p, pedal ganglia; pkm, pedal commissure; pr, prostate; ps, penial duct; rg, rhinophoral ganglia; rn, rhinophoral nerve; rs, seminal receptacle; ste, statocyst; udi, uterine duct; v, vestibulum; va, vagina; vg, vestibular gland. 17 December 2006, 6 m depth, Jeg. V. Padula. MNRJ 11973, three specimens, 35-40 mm alive, two dissected, Ilha do Papagaio, Cabo Frio, state of Rio de Janeiro, Brazil, 22°53’49"S, 41°59'2”W, 17 December 2006, 8 m depth, /eg. V. Padula. Distribution: Southeastern and southern coast of Brazil: states of Rio de Janeiro, Sao Paulo, and Santa Catarina (Dominguez et al., 2006; present study). External morphology: Living specimens ranging from 15 to 50 mm in length. Elongated body, slightly higher than wide. Adult background color dark blue to dark violet; body covered by longitudinal yellow lines (Figures 1D and 1E). Very young specimens with complete white mantle margin and having a complete yellow line on the central dorsum merged by two incomplete ones. Adults with three to nine dorsal longitudinal yellow lines. Lateral body walls and posterior dorsal region of the foot pale or dark blue with similar pattern of longitudinal yellow lines, sometimes not continuous. Genital opening surround- ed by a yellow ring in most (not all) specimens examined. Foot sole pale blue. Notum above eyes without pigmentation. Free notum border not undu- lated. Notum margin either completely yellow or white with small anterior and lateral yellow marks. Submar- ginal pale blue band with alternation of whitish and dark blue areas. Large MDFs (mantle dermal forma- tions) distributed along anterior and posterior notal border. Anterior foot margin bilabiate, upper lip with a yellow line, not notched. Oral tentacles short and digitiform, dark blue distally. Rhinophores retractile into moderately elevated sheaths. Dark blue rhino- phores perfoliate, with 20 lamellae in a 30-mm specimen; low gill sheath; longitudinal yellow lines ending at the margin, i.e., without encircling gills. Gill blue or violet, unipinnate or bipinnate. Gill rachis dark blue, with or without yellowish borders (Figures 1D and IE). Digestive system: Strong, muscular oral tube. Jaw rodlets stout to elongated, unicuspid, with curved tips (Figure 5A). Pharynx bulbous with posteroventrally projecting radula sac. Radula formulae 64 * 134.0.134 (ZSM_ 20040122, 20-mm fixed specimen) and 58 X 122.0.122 (MNRJ 11972, 35-mm living specimen). Narrow rachis, devoided of teeth. Lateral teeth hook- shaped, bicuspid; cusps slender, pointed, upper cusp generally longer than lower cusp, but of equal size in some teeth. Initial lateral teeth bicuspid; a triangular denticle can occur at base of the upper cusp (Fig- ure 5B). Some first lateral teeth having a single cusp. Midlateral teeth with 1—7 denticles under lower cusp; denticles may be blunt or pointed (Figure 5C). Outermost lateral teeth elongated with a blunt apical cusp; secondary blunt cusps present or absent (Fig- S. DaCosta et al., 2008 pa A Page 21 Figure 5. AHypselodoris picta lajensis (MNRJ 11972), SEM micrographs of hard parts. A, jaw rodlets; B, innermost radular teeth; C, midlateral teeth of the radula; D, outermost radular teeth. ure 5D). Salivary glands tubular, entering pharynx laterally to the thin-walled, tubular esophagus. Stom- ach and spherical caecum completely embedded in the holohepatic digestive gland. Intestine arising from terminal stomach towards the anterior region of the body, then curving to the right side and leading backwards to center of gill circle (Figure 6A). Circulatory system: Voluminous pericardium lying posteromedially in body cavity, above gonad and digestive gland. Two-chambered heart, longitudinally oriented; trapezoidal auricle, posterior to a muscular and spherical ventricle. Aorta divided into posterior aorta running into left side of viscera, and anterior aorta leading to head. Whitish blood gland, consisting of two flat lobes; posterior lobe partly covering salivary glands and distal reproductive system, anterior lobe very elongated and reaching the anterior region of body cavity. Syrinx leaving right pericardium ventrally to the atrium (Figure 6B). Nervous system: Comprising paired cerebropleural, pedal, buccal, gastroesophageal, and _ rhinophoral ganglia. Cerebral and pleural ganglia merged, larger than the other ganglia. Thickened rhinophoral nerves at base of rhinophoral ganglia. Rhinophoral ganglia located anterior to cerebropleural ganglia. Optic nerve short, situated laterally to cerebropleural ganglia. Statocysts placed laterally to cerebropleural ganglia, near pedal ganglia and beside eyes. Pedal ganglia ventral to, and slightly smaller than, the cerebropleural ganglia. Small buccal ganglia with short buccal commissure. Two gastroesophageal ganglia situated near buccal ganglia. Page 22 The Veliger, Vol. 51, No. 1 Figure 6. Hypselodoris picta lajensis, schematic anatomical drawings. A, digestive system; B, organization of major organ systems. Abbreviations: aa, anterior aorta; ao, anus opening; ap, posterior aorta; at, atrium; bgl, blood gland; ca, digestive caecum; dg, digestive gland; ey, eyes; gen, reproductive organs; go, gonopore; in, intestine; mo, mouth; oe, esophagus; ot, oral tube; ph, pharynx; r, radula; rhs, rhinophoral sheath; sg, salivary gland; st, stomach; ve, ventricle. Reproductive system: Gonad covers the digestive gland dorsally. Proximal gonoduct a long thin tube. Distal genital system triaulic (Figure 7). Hermaphroditic ampulla short and swollen. Distal gonoduct short. Oviduct thin, entering female gland mass after a short distance. Prostatic portion very long, with many loops attached to bursa copulatrix, situated above the ampulla. Distal vas deferens initially a thin winding tube, becoming a wider ejaculatory duct; without armature, slightly everted into elongate penial sheath. Male and female atria distally combined into common vestibule. Vagina a relatively wide tube. Large, spherical, thin-walled bursa (Figure 7). Receptaculum seminis club-shaped, size varying from one-half to one- third the diameter of the bursa, with a visible and curved stalk. Thin uterine duct splitting off the vagina distally to the receptacle stalk; uterine duct entering female gland mass close to the oviduct. Female gland mass well developed, consisting of several barely distinguishable winding glandular portions. Nidamen- tal duct showing conspicuous unilateral swelling; nidamental opening located posterior to the joint vestibule. Flat glandular lobe covering half of female gland mass; deferent duct hidden between nidamental opening and the vestibule (Figure 7). DISCUSSION Hypselodoris species from Brazil: Three species of Hypselodoris were previously known from Brazil (Marcus & Marcus, 1970; Troncoso et al., 1998; Dominguez et al., 2006b): (1) H. marci from the northeastern and southeastern coast; (2) H. sycilla from a single record on the northeastern coast; and (3) H. picta lajensis from the southeastern and southern coast. The taxonomy of these three species remains confusing, because the original descriptions and recent reports lack comprehensive descriptive data. Despite this, these three species can be easily distinguished from H. juliae sp. nov., which is characterized mainly by the combination of the following characteristics: a pale notum with many orange or yellow lines and iridescent S. DaCosta et al., 2008 \ \ fgm vg Figure 7. Hypselodoris picta lajensis, reproductive system. Abbreviations: am, ampulla; be, bursa copulatrix; ps, ejacu- latory duct; fgm, female gland mass; hd, hermaphroditic duct; nd, nidamental duct; no, nidamental duct opening; ov, oviduct; pr, prostate; ps, penial duct; rs, seminal receptacle; ud, uterine duct; v, vestibulum; va, vagina; vg, vestibular gland. blue patches, a relatively short prostate, and the absence of denticles on the lateral radular teeth. Hypselodoris marci is a yellowish species with a reticulate pattern of white, blue, and orange patches, and a grayish submarginal band with dark spots that borders the mantle (Marcus, 1971; Valdés et al., 2006). Hypselodoris marci possesses radular teeth with a series of small denticles (Marcus, 1971) and a long coiled prostate (personal observation). This species was originally described from northern Brazil and from Venezuela (Marcus, 1971). Later, it was reported from southeastern Brazil (Dominguez et al., 2006), Belize, and Honduras (Valdés et al., 2006). The assumption that specimens from different localities are conspecific was based mainly on external morphology. This may, however, lead to some taxonomic confusion, as did the supposed record of this species in the Caribbean Sea of Mexico (Ortea et al., 1996). The material originally illustrated by Ortea et al. (1996) corresponds to a different, recently described species, Hypselodoris olgae Ortea & Bacallado, 2007. Hypselodoris sycilla was originally described in 1890, from the Yucatan Peninsula, Caribbean Sea of Mexico, and remains a poorly known species. Valdés et al. (2006) suspected that H. sycilla could be synonymous with H. zebra from Bermuda. The single record of H. sycilla from Brazil refers to a single specimen (Marcus & Marcus, 1970), considered as a representative of H. picta by Ortea et al. (1996), but whose identity remains Page 23 unresolved. Like H. juliae sp. nov., H. sycilla is characterized by a pattern of longitudinal yellow lines. However, H. sycilla has a dark-blue notum (Bergh, 1890). Besides the differences in body color, H. juliae sp. nov. also differs from H. sycilla in radular morphology, the latter having small denticles on the radular teeth (Bergh, 1890), which are absent in H. juliae sp. nov. Hypselodoris picta lajensis is a dark-blue to violet species with well-spaced dorsal lines. The blue body sides also have yellow lines. Based on the original description and observations of additional material, Dominguez et al. (2006) separated H. picta lajensis from H. picta. The authors considered that H. lajensis has five dorsal yellow lines (against three lines in H. picta); uniformly deep blue-violet gills (against gill rachises with yellow lines in H. picta), and a deferent duct with a narrow preprostatic portion, which is absent in H. picta. However, our study of newly collected material broadened our perspective on morphological variation in Hypselodoris picta lajensis. Some specimens have up to nine dorsal yellow lines, rather than only five, and the gills have yellow lines on their rachises. Furthermore, the mantle margin of most of our specimens was completely yellow, not white with anterior and lateral marks, as originally described by Troncoso et al. (1998). Interestingly, this variation was observed between groups of specimens from two adjacent regions. All observed and collected specimens from Ilha Comprida, Cabo Frio (22°53'49"S, 41°59’2"W) have white mantle margins with yellow marks, whereas the majority of specimens observed or collected at Ilha do Papagaio, Cabo Frio (22°51'46’S, 41°56'32’W;, 5.7 km distant from Ilha Comprida) have yellow mantle margins. Differences between our material and that described by Troncoso et al. (1998) and Dominguez et al. (2006) also include radular features. Teeth were described as possessing upper cusps of approximately double the length of the lower cusps; an additional small denticle may be present on the lower cusp (Troncoso et al., 1998). In contrast, both cusps are nearly equally long in our material, and devoid of denticles on the lower cusp. Another difference was noted in the shape of the outermost lateral teeth. In our material, these teeth are a single plate without two major cusps, whereas bicuspid teeth were described for the type material (Troncoso et al., 1998). Sizes and proportions of allosperm receptacles also vary considerably in our material, but such variations probably reflect different stages of maturity and sexual activity. However, a differentiated, long, thin preprostatic portion, as illustrated originally (Troncoso et al., 1998, fig. 5), was not observed in the specimens that we examined. Furthermore, the postprostatic portion of our material is shorter and less convoluted. Because of these Page 24 variations, we prefer to maintain the subspecies rank of H. picta lajensis until a comprehensive revision of all material of H. picta, including detailed anatomical comparisons and also a molecular approach, can provide a better understanding of this group. Hypselodoris juliae sp. nov. clearly differs from H. picta lajensis in patterns of coloration; the former has a much clearer background color, and more and thinner longitudinal lines on the notum. The gill of H. juliae sp. nov. has dark rachises with orange margins, whereas H. picta lajensis has a dark-blue gill, with dark-blue rachises with or without yellowish margins. A white posterior mark is present on the rhinophores of H. juliae sp. nov. but absent in H. picta lajensis. Furthermore, the lateral teeth of H. juliae sp. nov. lack any denticles, whereas 1—7 denticles are present in H. lajensis. The prostate of H. picta lajensis is much more elongated and convoluted than the prostate of H. juliae sp. nov. Caribbean species similar to Hypselodoris juliae sp. nov.: Other species of Hypselodoris from the tropical western Atlantic resemble H. juliae sp. nov. with regard to background coloration and in having longitudinal yellow or orange stripes, i.e., H. ruthae, H. espinosai, and H. bayeri. Reaching up to approximately 30 and 20 mm in length, respectively, the former two species are considerably smaller than H. juliae sp. nov., have no more than half the number of notal stripes, and their free notal margins are not undulating as in H. juliae sp. nov. Furthermore, the lateral radular teeth of H. ruthae and H. espinosai have small denticles, which are absent in H. juliae sp. nov. (see Ortea et al., 1996). Hypselodoris bayeri is known from Florida, Cuba, Panama, Mexico, and Belize (Valdés et al., 2006). This species is blue with relatively broad yellow lines along the notum, body sides, and tail, which may be connected in a netlike pattern. The corresponding lines are more than twice as numerous and much narrower in H. juliae sp. nov. Submarginal rows of black dots along the notum and the foot are present in H. bayeri but absent in H. juliae sp. nov. The upper cusps of the lateral teeth are up to five times longer than the lower ones, and denticles are present in H. bayeri (see Ortea et al., 1996: fig. 65), whereas the cusps are equal in size or only slightly longer in H. juliae sp. nov. Caribbean specimens illustrated and listed as Hypselodoris sp. 3 by Valdés et al. (2006) externally resemble H. bayeri and, thus, differ from H. juliae sp. nov. in all the points mentioned above concerning external features. Hypselodoris juliae sp. nov. clearly differs from the nominal western Atlantic and other known species of Hypselodoris, and is thus established as a new species herein. Future studies of specimens of Hypselodoris sp. 1 of Valdés et al. (2006) and Hypselodoris sp. 4 of The Veliger, Vol. 51, No. 1 Rudman (2001-2003) will show whether or not they are conspecific with H. juliae sp. nov. Acknowledgements. We acknowledge Mark DaCosta (Bre- genz) for providing diving equipment and for help during field work. Dr. Peter Wirtz (Madeira Island) kindly sent us photographs of the specimen from Guarapari. Dr. Roland Melzer and Enrico Schwabe instructed us in preparing SEM micrographs, and Prof. Gerhard Haszprunar (all at ZSM) provided laboratory facilities and helpful advice. We also thank the CaboFrioSub staff, especially Carlos Alberto Filho, for the dive operations in Cabo Frio; Noémia Rodrigues, from the Laboratorio de Ultraestrutura Celular da Universidade Federal do Rio de Janeiro, for assistance with the SEM; Dr. Janet Reid and Dr. Juan Maceira for revising the English text. This study was supported by the GeoBioCenter'””, the German Research Foundation (DFG; SCHR 667-3) to MS, and the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq-Brasil) to VP. LITERATURE CITED BERGH, L. S. R. 1890. Report on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-1878), and in the Caribbean Sea (1879— 1880), by the U.S. Coast Survey steamer “Blake,” Lieut.- Commander C. D. Sigsbee, U.S.N., and Commander J. R. Bartlett, U.S.N., commanding. Report on the nudi- branchs. Bulletin of the Museum of Comparative Zoology 19:155-181. DacosTta, S., C. M. CUNHA, L. R. L. SIMONE & M. SCHRODL. 2007. Computerbased 3-dimensional recon- struction of major organ systems of a new aeolid nudibranch subspecies, Flabellina engeli lucianae, from Brazil (Gastropoda, Opisthobranchia). Journal of Mol- luscan Studies 73:339-353. DOMINGUEZ, M., F. J. GARCIA & J. TRONCOSO. 2006a. Some aspects of the family Chromodorididae (Opisthobranchia: Nudibranchia) from Brazil, with description of a new species. Scientia Marina 70(4):621—634. DOMINGUEZ, M., F. J. GARCIA & J. TRONCOSO. 2006b. A new species of Hoplodoris Bergh, 1880 (Gastropoda: Opistho- branchia: Nudibranchia) from the Atlantic Ocean. The Nautilus 120(4):150-155. GARCIA, F. J., J. S. TRONCOsO & M. DOMINGUEZ. 2002. New data on the benthic molluscs from the Archipelago of Fernando de Noronha (Brazil), with description of a new species of Aegires Lovén, 1844. Iberus 20:45—S6. GarCiA, F. J. & J. S. TRONcCoso. 2003. Two unknown species of Mollusca Gastropoda from the Archipelago Fernando de Noronha (Brazil), with description of a new species belonging to the genus Phidiana Gray, 1850 and a new record of Dendrodoris senegalensis Bouchet, 1975. Scientia Marina 67:159—166. GarciA, F. J. & J. S. TRONCOso. 2004. A new species of Anetarca Gosliner, 1991 (Gastropoda: Opisthobranchia: Facelinidae) from Western Atlantic Ocean. The Nautilus 118:139-143. Marcus, Er. & Ev. MARCUS. 1970. Opisthobranchs from Curacao and faunistically related regions. Studies on the Fauna of Curacao and other Caribbean Islands 122:1— 129. Marcus, Ev. 1970. Opisthobranchs from northern Brazil. Bulletin of Marine Science 20:922-951. Marcus, Ev. 1977. An annotaded checklist of the western Page 25 S. DaCosta et al., 2008 atlantic warm water Opisthobranchs. Journal of Mollus- can Studies Suppl. 4:1—23. ORTEA, J. & J. J. BACALLADO. 2007. Descripcion de una nueva especie de Hypselodoris Stimpson, 1855 (Mollusca: Nudibranchia: Chromodorididae) nombrada en home- nage a Olga Ucelay Sabina. Revista de la Academia Canaria de Ciencias 18(4):53—60. ORTEA, J., A. VALDES & J. C. GARCIA-GOMEZ. 1996. Revision de las especies atlanticas de la familia Chromo- dorididae (Mollusca: Nudibranchia) del grupo cromatico azul. Avicennia, suplemento 1:1—165. PADULA, V. & R. S. ABSALAO. 2005. Primeiro registro de Babakina festiva (Roller, 1972) (Mollusca: Nudibranchia) no Atlantico Sul. Biociéncias 13:99-101. PADULA, V. & F. N. SANTOS. 2006. Three new records of Nudibranchia (Mollusca, Gastropoda)—additions on the Brazilian biodiversity. Biociéncias 14(2):214—220. PoLa, M., J. L. CERVERA & T. M. GOSLINER. 2005. A new species of Tambja (Nudibranchia: Polyceridae: Nem- brothinae) from southern Brazil. Journal of the Marine Biological Association of the U.K. 85:979-984. Rios, E. C. 1994. Seashells of Brazil. Fundagao Universidade do Rio Grande, Centro de Ciencias do Mar, Museu Oceanografico, 331 pp. RUDMAN, W. B. 2001-2003. Hypselodoris sp. 4. Available at: http://www.seaslugforum.net/factsheet.cfm?base=hypssp4 (accessed in 10 January 2006). TRONCOSO, J. S., F. J. GARCIA & V. URGORRI. 1998. Anatomical data on rare Hypselodoris picta (Schultz, 1836) (Gastropoda: Doridacea) from the coast of Brazil, with description of a new subspecies. Bulletin of Marine Science 63:133-141. 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. WILSON, N. G. & R. C. WILLAN. 2007. Hypselodoris jacksoni, a new species from the southwestern Pacific Ocean (Nudibranchia: Chromodorididae), with a discussion on intraspecific variation in mantle glands in Chromodoris willani Rudman, 1982. Zootaxa 1549:29-42. The Veliger 51(1):26—42 (March 31, 2010) THE VELIGER _ © CMS, Inc., 2008 Diet and Feeding Habits of Octopus hubbsorum Berry, 1953, in the Central Mexican Pacific ERNESTO LOPEZ-URIARTE, EDUARDO RIOS-JARA Laboratorio de Ecosistemas Marinos y Acuicultura, CUCBA, Universidad de Guadalajara, Las Agujas, Nextipac, Zapopan, Jalisco. C.P. 45110. Tel: (52)33-37771156 (e-mail: ernlopez@cucba.udg.mx) AND MONICA ELIZABETH GONZALEZ-RODRIGUEZ Instituto de Acuicultura y Pesca, Secretaria de Desarrollo Rural, Gobierno del Estado de Jalisco Abstract. The diet and feeding habits of Octopus hubbsorum were analyzed using 226 individuals obtained from commercial artisan catches in the Central Mexican Pacific from July, 1999, to August, 2000. Organisms ranged from 43 mm to 230 mm in dorsal mantle length. The diet comprised 53 types in seven phyla; crustaceans, mollusks, and fishes were the main groups. In general, the crustaceans were dominant; in particular, species of brachyurans, carideans, and anomurans, with values of more than 40%, according to the index of occurrence and the indices of importance in weight and number. The diet is affected by sex, size, sexual maturity, and the season of the year. The females fed preferably on mollusks (gastropods and bivalves) and had a higher proportion of food in their stomachs than males, while the males fed mostly on crustaceans and members of the group “‘others.” The type of prey and its proportion also vary as the organism grows; the juveniles contained a lower number of prey species than the mature individuals. The males showed a clear tendency to increase the number of prey groups from warm to temperate environmental conditions. These results confirm that O. hubbsorum is an opportunistic predator. INTRODUCTION Octopus hubbsorum has a wide geographical distribu- tion, ranging from the central Gulf of California (28°55'N, 113°32'W) to the southern coast of Oaxaca, Mexico (16°10’N, 95°14’W) (Lopez-Uriarte et al., 2005). It constitutes practically half of the catch of the octopus fishery in the Mexican Pacific, totaling nearly 1000 tons per year (SAGARPA, 2002). During the last decade, this resource has occupied one of the first five places of the coastal fisheries of Jalisco (Rios- Jara et al., 2004). Octopuses are ferocious carnivores, feeding during the day or night on a wide variety of prey species, which are detected either by vision or by touch (Hanlon & Messenger, 1996). At all stages of development, octopuses are active predators, feeding mainly on crustaceans, mollusks, and fishes; but ophiuroids, polychaetes, chaetognaths, and siphono- phores constitute part of the diet of some species. The proportion of these types of food depends on the species, the sex, and the sexual maturity of the individuals (Nixon, 1987). Because of their opportu- nistic behavior, prey density also has an important effect on feeding; the octopods consume the most common prey available in their habitats (Wolterding, 1971; Hochberg & Couch, 1971; Hanlon, 1975; Van Heukelem, 1976; Ambrose & Nelson, 1983; Am- brose, 1984). However, mature females reduce their intake of food by up to 50% two weeks before spawning and for at least three weeks afterward; feeding is also reduced during the winter when temperatures are under 15°C (Borer, 1971). There have been no studies on the feeding behavior and diet of O. hubbsorum. The only previous report was made from observations of the shells of gastropods and bivalves found outside the caves and shelters of adult octopuses living in the shallow rocky areas of Bahia de Coastecomate, Mexico (19°13’47’N and 104°43'44"W) (Raymundo, 1995). The present study aims to describe, first, the diet of O. hubbsorum in the central Mexican Pacific through the analysis of stomach contents, and, second, the feeding dynamics using different indices to evaluate the possible effect of sex, size, stage of maturity, and the seasons of the year. This contribu- tion to the studies of the Mexican species of octopuses is the first detailed description of the feeding habits of this species. E. Lopez-Uriarte et al., 2008 Page 27 Wy 105° 00° u Pa legeae Islas Marietas _ Ne ( / Bahia Banderas if México JALISCO é 19°50 4 Punta Soledad ea Bahia Chamela Punta Pérula Bahia Tenacatita 90 Bahia Manzanillo Ao ne) Qo § 18°40" 4 Qs Punta Farallon Careyes Figure 1. MATERIALS AND METHODS Area of study: The littoral of Jalisco is located in the central Mexican Pacific (between 20°40’N and 18°58’N). This study was conducted in the central region of Jalisco, where Octopus hubbsorum is exploited commercially. This region extends approximately 38 km from Punta Soledad (19°36'47’N, 105°12’13”W) to Punta Farallon (19°23'22’"N, 105°02'17"W) (Figure 1). The coastline of this region is characterized by sandy beaches alternating with rocky shores and cliffs. In the shallow areas, there is a complex substratum of bedrock, boulders, rock rubble, and sandy bottoms, with small rocks inter- spersed with patches of sand covered with a layer of fine ‘sediment consisting largely of detritus particles. The more heterogeneous rocky bottoms have higher abun- dance and a greater variety of macroalgae, inverte- brates, and fishes. In some of these areas, several species of stony corals (Pocillopora spp. and Porites spp.) grow together, forming aggregations with a characteristic flora (macroalgae) and fauna (crustaceans, echino- derms, molluscs, polychaetes, gorgonians, and fishes). The region has warm—wet climate, with the rainy season occurring mostly during the summer. Temper- atures range from 32.3°C in September to 20.6°C in January (mean = 25.2°C) (Secretaria de Programacion y Presupuesto, 1981). The surface-water temperature of Bahia Chamela is higher from June to September (26.0-30.6°C) and lower from February to May (22.8— 26.6°C); the difference between the maximum value (July = 30.6°C) and the minimum value (February = 22.8°C) is approximately 8°C (Silva-Segundo et al., 2006). Cumulative monthly precipitation ranges be- tween 800 mm and 1500 mm, with the highest values from June to September and the lowest from February to April (Villalpando & Garcia, 1993). There is a mixed semidiurnal tidal cycle with two unequal high tides and two unequal low tides each day. The region is strongly influenced by tropical storms and cyclones during the warm-wet season. Coastal waters are relatively pro- Page 28 ductive and have the influence of three main surface currents: (1) the North Equatorial Countercurrent, with warm waters from the south (June—September); and (2) the Gulf of California Current (October— January) and (3) the California Current (February— May), both of which bring cooler waters from the north (Wyrtki, 1965, 1966). Therefore, there is well defined seasonality with three main seasons during the year: (1) a warm—wet season from June to September, which corresponds to the rainy period of the year, with warm surface-water temperatures higher than 26°C; (2) a warm—dry season from October to January, when the surface-water temperature is still warm and the rainfall is scarce or absent; and (3) a cool-dry season from February to May, when the surface temperature falls to approximately 22°C and there is low rainfall. Methods: A total of 562 individuals of Octopus hubbsorum were obtained from the commercial artisan catches made by local fishermen at three different localities of the central region of Jalisco. Time of capture was between 8:00 a.m. and 3:00 p.m. at depths shallower than 30 m. Collection of individuals began in July of 1999 and ended in August of 2000. The dorsal mantle length (DML), total body weight, and sex of each individual were first determined in the field, and then all individuals stored at —20°C. After thawing, the maturity stage was assigned in the laboratory using the scale of Guerra (1975) as revised by Cortez et al. (1995). To analyze the diet and feeding habits, a total of 226 specimens were used. The visceral mass of each specimen was separated (by dissecting the mantle) and weighed (+0.1 g). Also, the digestive tract was separated from the rest of the visceral mass and weighed (+0.01 g). The contents of the digestive tract was then separated and the prey items were counted and identified to the lowest possible taxon. The different types of prey were grouped according to taxonomic affinities, resulting in five putative groups: mollusks (including only gastropods and _ bivalves), cephalopods, crustaceans, teleosts (bonefishes) and other items. Several indices were used to describe the diet and to compare the items according to sex, size, stage of maturity, and the seasons of the year (Hyslop, 1980; Castro & Guerra, 1990; Sanchez and Obarti, 1993; Cortez et al., 1995): 1. Fullness Index FI =(Wsc/ Wr) « 100 Wsc = weight of the stomach contents. Wt = total weight of the specimen (Hyslop, 1980). The results of FY were grouped in levels according to the different degrees of filling of the stomachs, as suggested by Hernandez- Lopez (2000): FI = O (level I, empty stomach); 0 < FI = 0.3 (level II, almost The Veliger, Vol. 51, No. 1 empty stomach), 0.3 < FI < 1 (level III, half full stomach), FJ = 1 (level IV, full stomach). 2. Vacuity Index VI =(Es/Ts)*100 Es = number of empty stomachs, Ts = total number of stomachs analyzed (226). 3. Occurrence Index OJ = (We, /n SS Ge ) «100 Ne; = number of stomachs with the prey item i; n>S\y.' = total number of different prey species. 4. Importance in Weight Index JWI =(Wi/Wrt)*100 Wi = weight of each type of prey i, Wt = total weight of all prey items. 5. Importance in Number Index INI = (Ni/Nt)*100 Ni = number of individuals of each type of prey i, Nt = total number of individuals of all prey items. To analyze the influence of size on the diet and the feeding habits of O. hubbsorum, two groups were considered: (1) juveniles (60-100 mm DML) and (2) adults (>100 mm DML). Among the adults, three maturity groups were compared: immature, mature, and senescent (females) or discharged (males). Matur- ing specimens were included in the mature group. The seasons of the year considered for these analyses were (1) warm—wet season (June to September), (2) warm— dry season (October to January) and (3) cool-dry season (February to May). Comparisons of the indices (FI, VI, OI, IWIT, and INI) between groups were made by means of a chi- square test (Sokal & Rolf, 1969). The level of statistical significance used was a = 0.05. RESULTS Size and sex proportion of the octopus population: Figure 2 shows the size (dorsal mantle length, DML) and stage of maturity (immature, mature, and senes- cent) of 514 individuals of Octopus hubbsorum sampled from the commercial artisan catches in the central region of Jalisco. The DML ranged from 43 to 230 mm (mean = 110.43 mm + SD 42.78 mm). The sex proportion was 1.35:1 (296 females, 218 males). The larger sizes (DML) were recorded for the females (¢ = 3.29; P < 0.01), range 43 to 230 mm (mean = 111.36 + SD 26.33 mm) and the smaller sizes for the males, range 43 to 162 mm (mean = 104.49 + SD 21.49 mm). Composition and conservation of the prey items: A total of 53 different prey items were identified in the stomach content of the octopuses. These items included almost E. Lopez-Uriarte et al., 2008 Page 29 Oinmature 2mature senescent 2 35 SS ———eeEeEeEeEeEeEeEeaaaaaR7~(™%_wnDADESaESSEeeee——EEEEEEEEEEE NEN = — 30 25 N (=) Frecuency oO oa a | | =a Ill Bee See ee (NS 0) ee ee re ae ee SE eS 10 | | q fice sora === = oO oO Oo ie) oO jo) io) oO oO j=) o oO io) lo) i>) j=) i=) j=) i=) j=) Oo Oo oO fo) So = N ise) + wo © ~ foe) oO = N oO wt w ce) ~ 00 (op) oO WI N ise) + Ww) - Ss - - - - oa - - - N N N N N N Dorsal mantle length (mm) b 30 25 = se > 20 = (S) Cc ® =) oO © 15 LL 10 BO PTT TTT 0 10 20 0 0 0 0 70 Dorsal mantle length (mm) Figure 2. Page 30 The Veliger, Vol. 51, No. 1 Table | Composition of diet of Octopus hubbsorum. Ocurrence (NV) Weight (g) Number (J) total %o total Jo total Jo Crustacea 294 Si 225 46 2157 88 Estomatopoda 16 3% 22 1% Gonodactylidae Neogonadoctylus stanchi Pseudosquillidae Pseudosquilla adiastalta Caridea Wa 15% 192 8% Palaemonidae Pontonia sp. Alpheidae Alpheus malleator Alpheus lottini Alpheus sp.1 Synalpheus digueti Synalpheus nobilii Synalpheus sp. Anomura ; 14 3% 20 1% Porcellanidae Clastotoechus diffractus Pachycheles biocellatus Pachycheles panamensis Pachycheles sp. | Pachycheles sp. 2 Petrolisthes edwardsii Petrolisthes glasselli Petrolisthes haigae Petrolisthes sp. 1 Petrolisthes sp. 2 Brachyura 187 36% 1923 719% Mayjidae Mithrax denticulatus Mithrax sp. Thoe sulcata sulcata Teleophrys cristulipes Xanthidae Paractaea sulcata Microcassiope xantusii Xanthodius stimpsoni Liomera cintimana Platyactaea dovii Pilumnus gonzalensis Paraxanthias insculptus Mollusca 152 30% 124 26% 199 8% Cephalopoda 54 11% 86 18% 72 3% Octopodidae Octopus hubbsorum Octopus sp. 98 19% 38 8% Py) 5% Bivalvia Di 5% of 1% 36 1% Mytillidae Modiolus sp. Unidentified No. 1 Unidentified No. Unidentified No. 2 Unidentified No. 3 Gastropoda 71 14% 31 6% 91 4% Muricidae Unidentified No. 1 Unidentified No. 2 — Unidentified No. Unidentified No. 2 Pisces 41 8% 116 24% 47 2% Teleostei 41 8% 116 24% 47 2% Muraenidae Unidentified No. Unidentified No. 2 — — E. Lopez-Uriarte et al., 2008 Page 31 Table | Continued. Ocurrence (NV) Weight (g) Number (1) total Jo total %o total %o Others Di, 5% 19 4% 42 2% Annelida Polychaeta Unidentified No. 1 Unidentified No. 2 Chaetognatha Aphragmopohora Unidentified No. 1 Echinodermata Diadimastidae Unidentified No. 1 Unidentified No. 2 Equinometridae Unidentified No. 1 Unidentified No. 2 Ophiuroidea Ophiocomidae sp. Talophytas Padina sp. Total 514 100% 485 100% 2445 100% complete prey and fragments of several body structures in different stages of digestion. Only a small percentage (5%) was recorded as consisting of complete and freshly eaten organisms, while almost 95% of the material was fragmented or partially digested. Organ- isms from seven taxonomic groups were identified (Table 1): Thallophytes (macroalgae), Annelida, Ar- thropoda (subphylum Crustacea), Mollusca, Echino- dermata, Chaetognata, and Chordata (subdivision Teleostei). The taxonomic identification of crustaceans was based on the carapaces, rostra, dactyls, chelipeds, other appendages, and eggs in different stages of maturity. In the case of gastropod mollusks, the identification was made using the opercula and fragments of shells; in the case of bivalve mollusks, using fragments of the valves and the byssus. The octopod prey were recognized using body fragments (arms, beaks, and lenses) or the almost complete small juveniles; the egg masses of other octopuses were also present in the stomachs. Complete chaetognath individuals were found in the samples, but: only fragments of spines and body fragments of echinoderms, including their characteristic mouthparts. The polychaetes were identified using body fragments with multiple appendages. Recently caught fishes were almost complete, and other body parts (spines, bones, scales, and otoliths) were also present. Finally, the small pieces of macroalgae in the samples are probably an indication of the habitat or substratum where the prey were caught by the octopuses. Diet: The diet of O. hubbsorum is composed mainly of members of three groups: mollusks, crustaceans, and fishes. The crustaceans dominated in the samples, according to the frequency of their occurrence in the samples (57%), their weight (46%), and their number (88%). The brachyuran decapods occurred in 36% of the samples and represented 79% of the total number of prey. Other decapod crustaceans, the carideans, and anomurans, were less common in the samples (15% and 3%, respectively). The mollusks were the second most important group according to the frequency of their occurrence in the samples (30%), their weight (26%), and their number (8%). Cannibalism of O. hubbsorum was important in frequency (11%) and weight (18%), Table 2 Values of the Fullness Index (FI) for the females, males, and total individuals of Octopus hubbsorum. Content in the digestive tract: I (empty), II (almost empty), III (half full), IV (full). Females I II Ill IV I Juvenile 14.81 53.70 31.48 0 11.53 Adult 29.69 38.46 27.69 4.61 10.63 Males Total Il Ill IV I II Ill IV 53.84 34.61 0 13.20 53.77 33.01 0 70.21 19.14 0 21.42 51.78 24.10 2.67 Page 32 100 The Veliger, Vol. 51, No. 1 80 xo | — 607) © | Oo} TS = Bll 40 Bll i= = BV LL 20 70 100 110 120 130 140 Dorsal mantle length (mm) Figure 3. but it represented only 3% in number. Teleost fishes were the third most important taxonomic group, according to their occurrence in the samples (8%), weight (24%), and number (2%); the presence of juvenile moray eels (family Muraenidae) is probably unusual for O. hubbsorum. Finally, the prey items belonging to the phyla Talophyta, Annelida, Echino- dermata, and Chaetognata showed percentages lower than 5% in occurrence, weight, and number, and they were combined in the group ‘“‘others.’’ Most prey items of this group were not identified to species, and some were probably incidental, as in the case of the macroalgae. Fullness index (FI): The results indicate that O. hubbsorum feeds almost constantly in the area of study. Most octopuses analyzed (126 females and 100 males) contained prey in their digestive tracts with different degrees of fullness. According to the Fullness Index (FID), nearly 80% of the stomachs were in the categories II (almost empty) or IV (full) (Table 2; Figure 3). The amount of food in the stomachs was significantly different among females and males (X? = 4.73; df = 1; P < 0.05); the females recorded higher amounts than males. The total population showed a tendency to decrease the amount of food with the size of the individuals. This tendency was more evident among the females, more of which had stomachs in category II (almost empty). Category IV (full) was observed only in the stomachs of adult females. Although the values of FI were higher during the warm-—dry season, there were no significant differences between the seasons of the year (P > 0.05). Females showed higher FI values (>26%) than males during the period of study (X? = 8.72; df = 3; P < 0.03), except during the warm—dry season (P > 0.05) (Table 3). Vacuity index (VJ): Nearly 20% of the stomachs of O. hubbsorum were empty. There were no significant differences in the Vacuity Index (VJ) between females Table 3 Seasonal values of the Fullness Index (FI) for the females, males, and total individuals of Octopus hubbsorum. Content in the digestive tract: I (empty), II (almost empty), HI (half full), TV (full). Females I Il Il IV I Warm—wet 24.48 44.89 28.57 2.04 20 Warm-dry 7.69 53.84 38.46 0 12.5 Cool-dry 26.66 42.22 26.66 4.44 7.69 Males Total Il Ill IV I II Ill IV 70 10 0 23.18 52.17 23.19 1.45 45.83 41.66 0 10 50 40 0 67.30 25 0 16.66 55.21 26.04 2.08 E. Lopez-Uriarte et al., 2008 100 } Vacuity index (%) inmature mature senescent Figure 4. and males (P > 0.05). Figure 4 shows the tendency of the VI vary according to the maturity stage. There was no significant difference in the VJ between immature and mature individuals (P > 0.05). However, the proportion of empty stomachs increased to nearly 85% in the senescent females, and the value of the index was significantly different (X° = 16.39, P < 0.001). The values of the males and females were not significantly different in immature (P = 0.169) and mature (P = 0.056) individuals. On the other hand, there were significant differences in the proportion of empty stomachs observed between the seasons of the year. The warm—wet season showed the highest proportion of empty stomachs (37.82%), while the lowest was observed in the cool-dry season (X* = 6.55, P = 0.038) (Figure 5). Occurrence index (OJ), importance in weight index (JWI), and importance in number index (JNJ): The relative importance of each group of prey (crustaceans, mollusks, fishes, cephalopods, and “‘others’’) depended of the index in which it was expressed: OF, IWI or INI. The Kruskal-Wallis test indicated differences between juveniles (60-110 mm DML) and adults (>110 mm DML) in the indices of occurrence (OJ) (H = 41.61; P < 0.005), weight JWI (H = 31.96; P < 0.001), and number of prey (NJ) (H = 20.17; P < 0.028). The values of the three indices increased between the interval of 65mm and 110mm DML (Figure 6, 7, and 8), and decreased gradually toward the larger sizes. The JWI values showed differences in the feeding habits between males and females of Octopus hubb- sorum. The crustaceans and the group “others” were preferred by males, while the females preferred the mollusks (gastropods and bivalves) and fishes (Ta- ble 4). In the cases of cannibalism (cephalopod prey), there was no clear pattern of preference between sexes. The variety and proportion of new prey items in the diet increased with the size of the individuals. This was more evident in the case of crustacean prey, which Page 33 Warm-wet Cool-d an (37.82%) (22.69%) = , xR >, > KS RS Warm-dry (32.63%) Figure 5. declined in importance in the stomach contents as the size of the octopuses increased from 75 mm to 160 mm DML. Consequently, the importance of other prey (gastropods, bivalves, cephalopods, fishes, and “‘oth- ers’’) increased in these individuals, suggesting a change in the feeding habits with increasing size. This pattern was also observed in the JWI and INI (Figures 7 and 8). The Kruskal-Wallis tests indicate significant differences in the values of IWI and INI between the size intervals (7 = 41.74, P < 0.001; H = 20.17, P < 0.028, respectively). Dietary comparisons of the different groups of prey as a function of the size revealed that the mollusk prey were more important in juvenile individuals (<110 mm DML), while the cephalopod prey were more impor- tant in the adults (>110 mm DML) of both sexes (Table 4). The fishes were more important in the juvenile females and in the adult males. No differences were found in the crustaceans and the group “‘others”’ between juveniles and adults. The JWI showed higher values in the inmature individuals of both sexes for crustaceans and the group “others”? (Table 5); similar results were found in juveniles and adults (Table 4). The mollusks, cephalo- pods, and fishes were more important in the mature and senescent octopuses (Table 4b). The NIJ values showed significant differences between sexes only in the group “others”: the males had more prey items from this group (Table 6). All groups of prey had variations in the values of OI, IWI, and INI during the period of study (Figure 9). The males showed a more evident pattern; the three indices had a tendency to increase from the warm-—wet season to the cool-dry season. The cool— dry season not only showed the highest values of OJ, IWI, and INI, but also the highest proportion of cephalopods and teleost fishes. The crustaceans were dominant at all times, with values above 40% in both sexes. However, crustaceans increased their occurrence (JO) and weight (JW) in the samples during the warm—wet season (P < 0.05), but not their impor- Page 34 Ol crustaceans 100% ERS 90% 80% 70% oO e 60% ro) (0) = =) Q 50% — oO = 40% (ex ®O > 30% oO | LL 20% 10% 0% The Veliger, Vol. 51, No. 1 Nmolluscs “cephalopods &fishes others SQ ne n~n~©annygzemannnnnnn1“qmnmnon9nmimn0n0n9nnnnonny BAYOOOOOMOMOOMM yy SAAN ONOOMOOONOoonzy ANANSI SA a Saws / AAPA EA A nn~nsnww > b BOnnn~nnyfF OS 65 70 75 80 85 90 95 100 105 110 115 120 125 130 135 140 145 150 155 160 165 170 175 Dorsal mantle length (mm) Figure 6. tance in number (JN). The high proportion of crustaceans in the digestive tracts during the warm— wet season is remarkable, with percentages between 81% and 97% for the three indexes. During the warm-—dry season, the mollusks were significantly more important in the samples according to the OJ and JWI values (P < 0.05), but not according to the INT value. According to the OJ and the JWI, the females and males followed a similar pattern, since the participation of the five groups of prey increased from the warm—wet season to the temperate-dry season. However, this tendency was not clear with the INI. The participation of crustaceans was always important, with values between 40% and 90% of the INI. In general, mollusks (gastropods, bivalves, and cephalopods) were best represented during the warm period of the year (warm—wet and warm-—dry seasons). DISCUSSION AND CONCLUSIONS Previous studies on the diet of other species of Octopus report that they are active carnivores that feed mainly on crustaceans, mollusks, and fishes, while echino- derms, polychaetes, chaetognaths, and siphonophores form part of their diet in smaller proportions (Nixon, 1987; Guerra, 1978; Smale and Buchan, 1981; Am- brose, 1984; Cortez et al., 1995; Grubert et al., 1999). Additionally, these studies describe significant changes in the relative importance of the prey as a function of the sex, size, stage of maturity, and the season of the year. The diet and feeding habits of O. hubbsorum in the central Mexican Pacific is consistent with these descriptions. The method employed for assessing the diet is very important, since some procedures may overempha- size the importance of some kinds of prey relative to others. For example, the use of traps to asses the diet of O. vulgaris may result in a higher proportion of fishes relative to other prey, especially invertebrates, because octopuses and fishes were caught together in the traps (Hernandez-Lopez, 2000). Also, a procedure based on the collection of food debris around octopus middens would not record the consumption of fishes (Ambrose & Nelson, 1983; Ambrose, 1986; Mather, 1991). The only previous record on the diet of O. E. Lopez-Uriarte et al., 2008 100% 80% Oo oO Xf SSO 40% Weigth of prey (g) 20% 0% 65 70 75 80 85 90 95 100 105 110 115 120 125 130 135 140 145 150 155 160 165 170 175 Dorsal mantle length (mm) Figure 7. hubbsorum was based on this latter procedure (Raymundo, 1995), which indicated only remains of crustaceans and shells of gastropods and bivalves. The study of the content of the digestive tracts has been commonly used to study the diet of Octopus spp., and it is considered a reliable method because it gives evidence of the consumption of invertebrates and fishes with hard skeletons and shells that may be used for taxonomic identifications. A good evaluation of the diet of cephalopods should take into consideration a sample size that represents the different members of the natural populations, including juveniles, adults, and the senescent of both sexes. The sample size may be different according to the cephalopod species. Shchetinnikov (1986) estimated that approximately 20 stomachs per sample were enough to describe the diet of squid populations from the oceanic waters off Peru. Grubert et al. (1999) used a sample of 137 male ‘and female individuals to describe the diet and feeding strategy of Octopus moarum along the southeastern coast of Tasmania. In the Canary Islands, Hernandez-Lopez (2000) determined experi- mentally that a minimum sample size of 13 stomachs per month of Octopus vulgaris was sufficient to obtain 80% of the prey categories for this species. In the present study, a sample of 226 individuals of O. hubbsorum of both sexes from a wide range of sizes and different stages of maturity were analyzed during an annual cycle. The general coincidence in the feeding habits of O. hubbsorum with other species of Octopus and the ample variety of prey items found suggests that the methodology and the sample size were satisfactory. It is known that octopods are active hunters at night; they search for food mainly during the sunrise and sunset hours and make only short infrequent feeding trips during the day (Mather & O’Dor, 1991; Mather, 1991). The time of the day may also affect their feeding habits; for example, predation of O. vulgaris on fishes may change during the diurnal and nocturnal periods of the day (Nigmatullin & Osta- penko, 1976). Most octopods digest rapidly; in some species, the digestive process may last 14hr at temperate temperatures (Boucher-Rodoni et l., 1987); in the case of O. cyanea, individuals required 12 hr to complete the digestion at 30°C (Boucher- Rodoni, 1973). Our observations suggest that Octopus hubbsorum has nocturnal feeding behavior because most individuals were caught during the morning 100% 95% 90% > oO _ S 85% So 0 Same ® Qa = 3 80% Zz 75% 70% The Veliger, Vol. 51, No. 1 SSO PAR % SSO ra MA 65 70 75 80 85 90 95 100 105 110 115 120 125 130 135 140 145 150 155 160 165 170 175 180 Dorsal mantle length (mm) Figure 8. hours with food in their stomachs, indicating that the prey had likely been eaten recently or only a few hours before. However, since all individuals were obtained from commercial catches during a short time span (between 8:00 a.m. and 3:00 p.m.), there is no evidence of possible variations in the predatory behavior as a function of the time of the day. Future research should extend sampling to other periods of the day to obtain a better knowledge on the natural diet and the feeding habits of this species in the central Mexican Pacific. Octopus hubbsorum feeds on nearly 50 different prey species during the year. This is a relatively broad diet, compared with other species of Octopus; for example, 22 and 28 species were found in O. vulgaris (Ambrose & Nelson, 1983; Mather, 1991), 25 species were found in O. dofleini (Hartwick et al., 1981), and 12 species were found in O. maorum (Grubert et al., 1999). However, the dietary ranges of those other species are likely to be greater, since all of those studies were performed in relatively limited time spans (18 days to 8 months). The diversity of prey items found in O. hubbsorum is comparable with the 55 prey species found in a four- year study of O. bimaculatus (Ambrose, 1984), which was considered the practical limit of prey for that species. Similar groups of prey have been reported in natural populations of other species of Octopus, such as O. vulgaris (Nigmatullin & Ostapenko, 1976; Guerra, 1978; Hatanaka, 1979; Smale & Buchan, 1981; Sanchez & Obarti, 1993; Hernandez-Lopez, 2000), O. bimacu- latus (Ambrose, 1984), O. mimus (Cortez et al., 1995) and O. maorum (Grubert et al., 1999). All of those species feed mainly on crustaceans, mollusks, and teleost fishes, although their relative importance in the diet varies as a function of the species. Since cephalopods are opportunistic predators, they should consume different prey according to their availability. Therefore, the area in which the species live should affect their diet (Nixon, 1987; Hartwick et al., 1981; Ambrose, 1984). Thus, the same species may have differences in the types of prey consumed as a function of the distribution of the populations. This is evident in the case of the populations of O. vulgaris from the North Atlantic coast of Africa (Guerra, 1978) and the Mediterranean Sea (Sanchez & Obarti, 1993). The variety and availability of the prey is, in turn, generally related to the complexity of the habitat. Octopus hubbsorum is the target species in the octopus fisheries of the Mexican Pacific (Aguilar- Chavez, 1995; Rios-Jara et al., 2001). The fishermen Page 37 100°0> 100°0> 100°0> 100°0> 100°0> :d 1168 3°08 £96 etl 1 vol (pb = 13) x OL O8S Sol ILS 898 OCL 86S 08S 868 MOOT [PIOL es) 1k 8e 1 19'p eK cs'0 su 08°0 119 ok cs'0 CLT xx 08°0 JIYWIO bell ek ISO LVL su OO'L ek ce Cl vo Tl ok 00°L IS'0 xx CECI Toysoofe L 88 El su Loel 9L'OI eK PS 87 1k tc6 88'el Ok PS 8c LoEl * t£6 vpodojeydaD Sc Il ee £09 Iv9e x 8L Cl 1k 00°61 ST Il su 8L Tl £09 ek 1061 BosnT[OW 08°9S ee 6L°89 COI Ok els 4k CS°8S 08°9S * ells 6L°89 xe CS 8S vooRIsNID ane on} BUI] yUdDS9UIS oinj}eyy ony eu] oO fo) oO 6 Kald soley So[BWId,] ainjeyy din} eu] ‘quroytusis ATYSIY = x» ‘VUROIIUBIS = , “Wunsosqqny sndojIQ JO S[enplAIpUt dy} JO AjLINyeUL JO o3vIS pu soxas UIIMIOq (TAI) XOPU] IYBIOAA Ul 9oUvIIOdWY OY) Jo suostIeduIOD § IGeL 100°0> 100°0> 100°0> 100°0> 100°0> ‘d a4 v'6L 9°STI 6°98 VI9tl (p = 13) 65 DML) adult and senescent individuals of O. hubbsorum increased the proportion of cephalopods, fishes, and the group “others” in their diet, and they reduced the consumption of crustaceans and mollusks. These individuals ingested proportionally more types of prey than the juveniles. This is a common feature of cephalopods (Boucher-Rodoni et al., 1987; Hanlon & Messenger, 1996), and it has also been documented in other species of Octopus (Nigmatullin & Ostapenko, 1976; Guerra, 1978; Castro & Guerra, 1990; Cortez et al.,1995). Other reports on feeding habits of cephalopods suggest that there is not a large difference between the diet of juveniles and that of adults among the coastal species of octopuses (Boucaud-Camou & Boucher-Rodoni, 1983; Bouche-Rodoni et al., 1987). E. Lopez-Uniarte et al., 2008 Females INI (%) Females Warm-wet . Z 0 10 20 30 40 50 60 70 80 90 100 IWI (%) Females Ol (%) 0 10 20 30 4 0 10 20 30 40 50 ee: [= a = ae = 30 4 50 60 70 80 90 100 INI (%) Males Wanye ta Pa ag 0 70 80 90 100 IWI (%) Males pe Ol (%) Figure 9. Hernandez-Lopez (2000) reported some overlap in the diet among juveniles and adults of O. vulgaris. In the case of O. vulgaris, Guerra (1978) reported a change in the diet with respect to a depth gradient, particularly in the importance of the species of crustaceans consumed. There were also significant differences in the diet between adult males and females of O. hubbsorum, which were probably related to the higher reproduc- tive costs of females. A higher proportion of food by weight in adult females than in adult males of O. mimus has been associated with differences in the Page 40 nutritional needs of the two sexes (Cortez et al., 1995). Female octopuses require more energy for reproduction than males (O’Dor & Wells, 1978). In O. vulgaris, the high deposition of lipids in the yolk has been considered as a limiting factor for egg production by adult females (O’Dor et al., 1984). This may be true also of O. hubbsorum on the coast of Jalisco. Octopus hubbsorum had a higher proportion of empty stomachs in mature and senescent females than in males of the same stages. A_ similar condition has been reported in other species, such as O. mimus, from the coast of Chile (Cortez et al., 1995); the mature females of this species have less opportunity of catching prey (Mangold, 1987). This effect has been attributed to an inhibition of the appetite due to hormonal changes during this stage in the life cycle (Wodinsky, 1978). Also, the senescent females of O. hubbsorum ingested propor- tionally more items of the group “‘others” than did the senescent males. Cortez et al. (1995) suggested that the increased ingestion of the group “‘others” among the senescent females of O. mimus may be related to a reduction in their feeding activity associated with their need to remain close to the eggs while brooding, so that they tend to ingest more of the food available around the octopus middens, including those small and not very motile invertebrates of the group “others.” During the annual cycle, the composition of the diet of O. hubbsorum did not vary very much; the crustaceans were dominant at all times, with values above 40% in both sexes. The other groups of prey had variations in their occurrence, weight, and number, depending on the sex and the season of the year. The males did not feed on cephalopods and fishes during the warm—wet season, while females did not feed on the group “others” during the warm—dry season. However, variations in the proportions of prey items were evident among the seasons of the year. This behavior is similar to that reported for O. mimus, in which the main prey, the fishes, decline in proportion during fall and winter as the importance of other prey, such as crustaceans, increases (Cortez et al., 1995). There is a similar change in the diet of O. vulgaris, in which fishes and octopods_ that constitute the main items of prey during winter and spring are replaced by crustaceans at the beginning of summer. In summary, the diet and feeding habits of O. hubbsorum in the central Mexican Pacific are consistent with previous descriptions for other Octopus spp. Species-specific related behavior could be the cause of some differences, but these variations may also be related to the regional distribution of the populations studied. The results of the present study The Veliger, Vol. 51, No. 1 indicate that O. hubbsorum is an _ opportunistic predator that feeds during the night on a wide variety of prey. More detailed studies using different sampling methodologies at different times of the day and night are needed to allow us to learn more about the natural diet and the feeding habits of this species. Acknowledgments. Our thanks to local fishermen at Careyes, Punta Pérula, and Chamela for providing the individuals of Octopus hubbsorum used in this study. The professors and students of the Laboratorio de Ecosistemas Marinos y Acuicultura helped during data collection in the field. This work had the financial support of SMORELOS-CONACyT (Project No. 1998-03-06-21) and Universidad de Guadalajara, México. LITERATURE CITED AGUILAR-CHAVEZ, S. G. 1995. Estudio biolégico-pesquero del pulpo Octopus sp. (Cephalopoda: Octopoda) de la costa sur del estado de Jalisco. Tesis profesional. Centro Universitario de Ciencias Biologicas y Agropecuarias. Universidad de Guadalajara: Guadalajara, México. AMBROSE, R. F. 1984. Food preference, prey availability, and the diet of Octopus bimaculatus Verrill. 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University of Hawaii, Honolulu. WYRTKI, K. 1965. Suface currents of the eastern tropical Pacific Ocean. Bulletin of the Inter-American Tropical Tuna Commission 5:271—291. Page 42 WyRTKI, K. 1966. Oceanography of the Eastern Ecuatorial Pacific Ocean. Oceanography and Marine Biology Annual Review 4:33-68. WODINSKY, J. 1978. Feeding behaviour of broody female Octopus vulgaris. Animal Behavior 26:803-813. WOLTERDING, M. R. 1971. The rearing and maintenance of Octopus briareus in the laboratory, with aspects of their The Veliger, Vol. 51, No. 1 behaviour and biology. Master’s Thesis. University of Miami, Coral Grove, Florida. YANEZ-RIVERA, J. L. 1989. Estudio ecologico de las comunidades de gasterOpodos macroscopicos de algunas playas rocosas de la costa del estado de Jalisco, México. Tesis de Licenciatura. Facultad de Ciencias, Universidad de Guadalajara. Guadalajara, México. THE VELIGER The Veliger 51(1):43-62 (March 31, 2010) © CMS, Inc., 2008 Sacoglossan Opisthobranchs on Northwestern Pacific Shores: Stiliger berghi Baba, 1937, and Elysia sp. on Filamentous Red Algae CYNTHIA D. TROWBRIDGE* Oregon Institute of Marine Biology, University of Oregon, Charleston, Oregon, USA YOSHIAKI J. HIRANO Department of Biology, Graduate School, Chiba University, Japan and Marine Biosystems Research Center, Faculty of Science, Chiba University, Japan AND YAYOI M. HIRANO Marine Biosystems Research Center, Faculty of Science, Chiba University, Japan Abstract. At least 20 species of sacoglossan opisthobranchs worldwide feed on delicately branching red algae; these species include members of three genera (Hermaea Lovén, 1844; Stiliger Ehrenbergh, 1831; and Elysia Risso, 1818) in three sacoglossan families. The algal hosts include members of three algal families and at least ten algal genera in the order Ceramiales. We studied two sacoglossan species that feed on filamentous red algae: (1) the temperate to boreal Stiliger berghi Baba, 1937, on wave-sheltered shores of Honshu and Hokkaido, Japan, and (2) the subtropical to tropical Elysia sp. on moderately wave-exposed shores of Okinawajima and Honshu, Japan. Preference experiments demonstrated that S. berghi prefers to associate with the alga Dasya when given pairwise algal choices but readily consumes members of several algal genera and exhibits no preferences between algal life- history phases (diploid tetrasporophytes vs. haploid female gametophytes). Elysia sp. is a small sacoglossan that consumes uniseriate and polysiphonous red algae. Given the small size and seasonally abundant populations of organisms that feed on red algae, we predict that these sacoglossans and their ecological analogs on other shores may have an unexpectedly important role in consuming and/or fragmenting ceramialean red algae. Given the known propensity of these algae to be dispersed by international shipping and oyster mariculture, careful malacological consideration should be given to the potential of sacoglossans to be inadvertent “hitchhikers” on a global scale. INTRODUCTION overlooked. Furthermore, surprisingly little malacolog- ical attention has been focused on the possibility that many of the newly reported Hermaea spp. may not be indigenous to the regions from which they have been described. The taxonomy and ecology of these specialized herbivores on native and introduced ceramialean hosts in different areas of the world merits investigation. Throughout temperate and tropical areas of the world, there are numerous sacoglossan opisthobranchs that feed on species of Ceramiales; these sacoglossans belong to at least three sacoglossan families and three genera: Hermaea Lovén, 1844; Stiliger Ehrenbergh, 1831; and Elysia Risso, 1818 (Table 1). Our objectives of this report are threefold: (1) to review and compare the sacoglossan opisthobranchs on ceramialean red algae worldwide, (2) to evaluate the * Mailing address: PO Box 1995, Newport, OR 97365 USA; similarities and differences of the algal hosts, and (3) to e-mail: cdt@uoregon.edu describe the reproductive features and feeding ecology Marine red algae have been extensively introduced around the world by the fouling of ship hulls, the transport and relaying of oysters, and the introduction of epiphytized host plants. Of the macroalgae currently recognized as introduced (reviewed by Trowbridge, 2006), members of the red algal order Ceramiales predominate. The dramatic proliferation of nonindig- enous red algae (e.g., Womersleyella setacea covering the benthos in the Mediterranean and “‘Heterosiphonia japonica’ on Atlantic European shores) is widely recognized among phycologists and invasion biologists but not among malacologists. 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D. Trowbridge et al., 2008 jo Joquinyy jo Jo}oWIeIq = wnuwiIxey| ‘ponunuod T 219% L Page 46 of two Japanese species (Stiliger berghi Baba, 1937, and Elysia sp.) to enhance future identification of these species and to stimulate additional records within the North Pacific region. SACOGLOSSAN REVIEW Within the genus Hermaea, the most abundant species (in terms of population density) is Hermaea bifida (Montagu, 1815); despite its frequency, the species’ preference for native vs. introduced red algae on European shores has been overlooked. The uncommon to rare congeners include H. variopicta (Costa, 1869), which is considered by some authors to belong to genus Hermaeopsis A. Costa, 1869; H. paucicirra Pruvot-Fol, 1953, on NE Atlantic shores; H. cruciata Gould, 1870, on NW Atlantic shores; H. oliviae (MacFarland, 1966) and H. hillae Marcus & Marcus, 1967, on NE Pacific shores; H. wrangeliae (Ichikawa, 1993) on NW Pacific shores; H. noto (Baba, 1959) on NW Pacific shores; and H. evelinemarcusae Jensen, 1993, on Australian shores. [Note that Ichikawa reported her species as Aplysiopsis wrangeliae, but that genus contains only green-algal feeders, not red-algal ones; A. oliviae was moved into Hermaea (Rudman, 2000, with a 2001 update; Behrens & Hermosillo, 2005), based largely on diet. The first author (CDT) considers A. wrangeliae to belong to the genus Hermaea, though the absence of a voucher and the lack of descriptions of radular teeth and the reproductive system hinders confirmation of the genus.] Numerous additional Hermaea spp. have been record- ed: H. boucheti Cervera et al., 1991 (now considered to be H. bifida by Cervera et al., 2004), and H. ghanensis Caballer et al., 2006, on NE Atlantic shores; H. nautica Caballer et al., 2007, on NW Atlantic shores; H. coirala Marcus, 1955, on SW Atlantic shores (considered to be conspecific with H. cruciata by many colleagues); H. zosterae (Baba, 1959) on Japanese shores (considered to be conspecific with H. noto by some colleagues; see Rudman, 2002); and congeners in the Indo-Pacific region and Southern Ocean (e.g., Carlson & Hoff, 2003; Burn, 2006; Sea Slug Forum, www.seaslugforum.net). These species are often assumed to feed on red algae, although observational evidence is still frequently lacking. There are at least two Stiliger species that feed selectively on filamentous red algae: Stiliger fuscovitta- tus Lance, 1962, is an abundant, though insufficiently studied, sacoglossan on NE Pacific shores (Lance, 1962; Case, 1972; Trowbridge, 2002) and S. berghi Baba, 1937, on NW Pacific shores (Table 1). The NE Pacific specimens of S. fuscovittatus belong to the genus Stiliger, not to Ercolania: the teeth are blade-shaped, not sabot-shaped, and the species eats red algae, not green algae (characteristic of Ercolania spp.). [The true identity of “Ercolania fuscovitatta’” recorded from The Veliger, Vol. 51, No. 1 Florida and the Gulf of Mexico is presently unclear, as the radular tooth shape has never been reported; see discussion on Sea Slug Forum]. The NW Pacific S. berghi was described from 15 specimens collected in 1935 from Tomioka, Amakusa, Japan (Baba, 1937); the species was subsequently reported from Osaka Bay, Inland Sea of Seto, and Toyama Bay (Inaba, 1958, 1962; Baba, 1959; Hamatani, 1961). This species has rarely been reported, presumably owing to its small size, cryptic coloration, and patchy distribution. It has not been included in any of the recent popular books on Japanese opisthobranchs (Suzuki, 2000; Nakano, 2004; Ono, 2004) or a recent Japanese sacoglossan account (Hamatani, 2000). The only literature report for this species outside of Japan is from Russian shores—Peter the Great Bay, Sea of Japan (Adrianov & Kussakin, 1998). Not only is there distributional uncertainty but there is also taxonomic uncertainty. Baba & Hamatani (1970, p. 202) suggested that S. berghi may “‘belong to another genus, not yet defined.” Finally, at least four Elysia spp. consume red algae (Table 1). Elysia viridis (Montagu, 1804) on European shores feeds not only on four genera of green algae but also on four genera of red algae, including native and introduced species. The Australian Elysia cf. furva- cauda consumes green algae in some months and unidentified red algal epiphytes during other times (Brandley, 1984). On Japanese shores, Elysia abei/ amakusana feeds on the red alga Griffithsia as well as on green algae (Bryopsis, Cladophora, Chaetomorpha, etc.; Y.J. Hirano & Y.M. Hirano, unpublished observations). Originally described as separate species, E. abei Baba, 1955, and E. amakusana Baba, 1955, are currently presumed to be conspecific by many mala- cologists (see www.seaslugforum.net). Finally, on Okinawan shores, there is a small Elysia sp. that feeds on uniseriate and polysiphonous filaments of red algae. One specimen of this species has also been collected from Shimoda, Izu Peninsula, Honshu. Our unde- scribed Elysia sp. may be conspecific with Elysia sp. 2 from Hachijo-jima, Honshu (Nakano, 2004), Elysia sp. 5 from the Ryukyu Islands (Ono, 2004), or an Elysia sp. in the Philippines (T.M. Gosliner, personal communication). Until the internal anatomy of these specimens has been characterized, we will limit our discussion to our Elysia sp. ALGAL HOST REVIEW All of the sacoglossans that associate with red algae use hosts within the order Ceramiales; there are no confirmed reports of hosts from other algal orders (Table 2). Intriguingly, the hosts belong to three algal families: the high-diversity Rhodomelaceae and Cera- miaceae as well as the comparatively lower-diversity Dasyaceae (Table 2). As these families are distinguished C. D. Trowbridge et al., 2008 Page 47 Table 2 Classification of described hosts of sacoglossans that feed on red algae, based on currently recognized names in AlgaeBase (Guiry et al., 2006). The number of algal genera that are used as hosts are shown relative to the number of genera described in each algal family (based on Guiry et al., 2006). References Baba (1937, 1959); MacFarland (1966); this report Van Bragt (2004); Lance (pers. comm. to CDT); Hansen (pers. comm. to CDT) Dasysiphonia Van Bragt (2004) Heterosiphonia Thompson (1976) Orders Families | Number of genera Genera Ceramiales Rhodomelacae 1/137 (0.7%) = -Polysiphonia Dasyaceae 3/18 (16.7%) Dasya Ceramiaceae 6/142 (4.2%) Ceramium Griffithsia Halurus Bornetia Wrangelia this report Robertson (1868) '; Trowbridge (unpubl. obs.); this report Wright (1859) 7; Duerden (1896); Miller (1958); Kremer & Schmitz (1976)°; Trowbridge (unpubl. obs.)* Cornet & Marche-Marchad (1951) Ichikawa (1993) Callithamnion Lance (1962), Trowbridge (2002) 'G. corallina is considered to be G. corallinoides. 7G. setacea and 3G. flosculosa are considered to be Halurus flosculosus. primarily on reproductive attributes, what determines which families and genera within these families are appropriate hosts for sacoglossans? All host genera are either monosiphonous (with a single row of cells attached end-to-end) or polysiphonous (with a central series of cells surrounded by pericentral cells of identical height) with large cells. Polysiphonous hosts are frequently assigned to the genus Polysiphonia (Tables 1 and 2). However, given that the taxonomic status of the high- diversity genus has been historically in constant flux and was recently subdivided into multiple polysiphonous clades (e.g., Womersleyella, Neosiphonia, Polysiphonia, etc.; see Choi et al., 2001), the number of polysiphonous host genera recognized by malacologists is undoubtedly an underestimate. Cortication—the covering of the algal axis with a layer of small cells—occurs in a few host genera (e.g., Ceramium, Dasya), but most host genera lack cortication. Ceramium exhibits regions of cortication and regions without, thus providing saco- glossans access to small cells and large cells, respective- ly. Some genera, such as Dasya, have highly corticated main axes but monosiphonous branchlets. What is not clear, however, is why many red-algal genera, with seemingly appropriate morphology for sacoglossan feeding, are not reported as hosts. To what extent are the results in Table 2 a reflection of actual host use versus an artifact of challenging phycological identifi- cations by malacologists? We predict that future collaboration between sacoglossan researchers and taxonomists of red algae will reveal substantially greater host diversity than is currently recognized. METHODS Study regions: The present investigation was conducted primarily at three regions in Japan: (1) the Oshoro Marine Station, Hokkaido University; (2) the Misaki Marine Biological Station (MMBS), on the Miura Peninsula, Honshu, University of Tokyo; and (3) Okinawajima, the main island of Okinawa. The Oshoro Marine Station is on the southwestern side of Hokkaido on the Sea of Japan (Figure 1A); collections were made at several sites within the wave-sheltered bay. MMBS is on the Sagami Bay shores of the Miura Peninsula on the Pacific coast of Honshu; collections were made from Araihama and Moroiso Bay at MMBS (Figure 1B). Finally, we collected sacoglossans from three sites (Zanpa-misaki, Sobe, and Sunabe) on the southwestern coast of Okinawajima (Figure 1C). Collections and experiments: Our specific field collec- tions were as follows. In May 2000, we collected Codium fragile ssp. fragile (covered with epiphytes of the red alga Polysiphonia sp. and the sacoglossan Stiliger berghi) from ropes suspended within Moroiso Bay at MMBS. We examined the single egg mass produced by S. berghi in the laboratory (field-collected masses had already undergone several cell cleavages). We measured the maximum and minimum length of 10 uncleaved ova and capsules within the egg mass. When veliger larvae hatched from the spawn mass, we measured the maximum shell length of 10 veligers. In July 2001, we collected >100 Stiliger berghi on free-living red algae in various areas on the eastern shore of Oshoro Bay, Hokkaido (Figure 1A). We conducted short-term feeding-preference experiments at the Oshoro Marine Station, Hokkaido University. We offered individual specimens of S. berghi pairwise choices of three algal hosts: Dasya sessilis, Polysiphonia sp., and Ceramium sp. We monitored the number of specimens on each alga periodically throughout a one- day period and compared slug choices with Pearson’s Pacific Ocean Sea of Okhotsk Pacific Ocean The Veliger, Vol. 51, No. 1 ik Miura Peninsula | Hayama — a Sagami Bay Uraga Channel Araihama & Moroiso Bay A, Figure 1. Map of three major study areas in Japan from north to south: (A) Oshoro Bay, Hokkaido; (B) Miura Peninsula, Honshu; and (C) Okinawajima, Okinawa. chi-square test and Fisher’s exact test on the categorical data. In spring 2004, 2005, and 2006, we collected epiphytic and free-living Polysiphonia spp. from ropes suspended in Moroiso Bay. We also made collections of Zostera from Moroiso Bay (with red algal epiphytes) and ceramialean red algae from the adjacent rocky shore, Araihama, at MMBS. In March 2004, we hatched veligers to remeasure shell length. Furthermore, we conducted another feeding experiment to determine whether Stiliger berghi exhibited feeding preferences for thalli of different life- history phases. Red algae belonging to Ceramiales typically have a complex life history with haploid female and male thalli (gametophytes) coexisting with morphologically similar diploid thalli (tetrasporo- C. D. Trowbridge et al., 2008 phytes). Herbivores that feed on these algae may exhibit preferences between the isomorphic haploid and diploid phases. In March 2004, we conducted a pairwise-choice experiment with female gametophytes and tetrasporophytes of Polysiphonia harveyana. Prof. Chris Maggs (Queens University, Belfast) kindly identified the species on morphological and molecular grounds. The logistical details were similar to those of the experiment above. In April 2008, we collected 8 specimens of S. berghi from Polysiphonia (sensu lato) spp. and | from Ceramium sp. on the floating docks at Hota on the eastern shore of Sagami Bay, Honshu. The specimens were held in separate small containers filled with seawater and offered the red alga Griffithsia japonica. Observations were made a few times a day for 6 days to evaluate whether the sacoglossan could consume Griffithsia. Finally, we made seven visits to Okinawajima between July 2002 and April 2006. We collected ceramialean red algae primarily at Sunabe, Chatan, on the reef crest and at Zanpa-misaki on the reef platform. Smaller collections were also made in reef- crest and reef-lagoon areas of Sobe. Small, cryptic sacoglossans could not be observed directly on the shore. Therefore, specimens of Elysia sp. were sampled by collecting red algae, holding them in shallow trays with seawater for | to 2 days, and then hand-picking the slugs from the algal surfaces or the air—water surface. Because of the physical conditions (moderately wave-exposed habitats) and the ecological ones (intense herbivory), the red-algal thalli were typically <1 cm long and could be identified only to genus. Specimens of Elysia sp. were held in the laboratory in containers with different algal genera. Because of low sample size, formal experiments were not conducted, but feeding damage on different algal genera was recorded when observed. The single egg mass produced in the laboratory was collected; the diameters of ova and capsules were measured. RESULTS Stiliger berghi External features: The simple rhinophores of Stiliger berghi had a distinctive brown stripe encircling the upper end (Figure 2A—C). There was scattered purple- brown pigment on the epidermis on the dorsal surface; the sole of the foot was white although the pigment was present around the margin. This species had a very pronounced C-shaped, brownish-purple esophageal pouch (Figure 2B, C). The ingested red algal chloro- plasts were visible in the digestive tract, particularly in specimens that had recently fed. There were two longitudinal tubes that extended branches into the cerata but not into the tail. The anus was small and dorso-anteriorly situated. Page 49 The species grew to 10 mm long and 11 mg in wet weight, but most specimens were much _ smaller (Figure 3A—C). Of 45 specimens examined in April 2006 (Figure 3D), Stiliger berghi had 4 to 11 cerata per side (mean = 15 cerata total, maximum = 21); however, some large specimens collected on 4 March 2004 had 15 cerata per side. The species had pointed radular teeth with an unusual knob-like tip (Figure 4A—C). The ventral side of each tooth had a concavity, and the tip was blade- like with small denticulations on the central, keel-like edge. An extremely unusual feature was that the denticulations were on both the ventral and the dorsal edges of each tooth tip (Figure 4C). Based on 17 specimens (1—3 mm long) collected in July 2001 from Oshoro Bay, the number of ascending teeth varied from 6 to 8; the number of descending teeth varied from 17 to 23, including the three preradular teeth. The length of the leading tooth varied from 80 to 135 um; the base width varied from 25 to 40 um. Our observations of tooth shape, size, and number are consistent with the species description, with two rather minor exceptions. (1) Baba (1937) mentioned the highly distinctive teeth were smooth, based on light micros- copy; with currently available, high-resolution micros- copy, the denticulation is readily apparent. (2) The original description states there were five teeth in the descending row and 19 teeth in the ascending row; we respectfully suggest that Baba inadvertently switched the terms ascending and descending. Reproduction and development: Stiliger berghi copulated frequently in the laboratory. Slugs started by vigor- ously pressing the anterior part of the head together with that of a conspecific. The slugs then aligned themselves right eye to right eye. Copulation took several minutes and was reciprocal; the short penis lacked a stylet. The slugs copulated sequentially with many different slugs. The egg masses of S. berghi were oval and were deposited among the branches of the polysiphonous red algae. The slightly oval ova averaged 62 X 68 um in diameter, and capsules were 104 X 136 um. When veliger larvae hatched from the spawn mass in May 2000, initial shell length averaged 116 wm. In March 2004, recently hatched veligers averaged 114 um in shell length (n = 5) and had type I shells (sensu Thompson, 1961). Feeding ecology: We found several algal hosts and observed feeding activities of Stiliger berghi. In May 2000, some of the subtidal thalli of Codium fragile ssp. fragile that we collected from Moroiso Bay had the epiphytic red alga Polysiphonia sp. We found 33 individuals of S. berghi on five thalli of C. fragile with this epiphyte. Cell size of the epiphyte on the main axes was about 200 um wide and 300-400 um long; on Page 50 The Veliger, Vol. 51, No. 1 Figure 2. (A) Oblique right view, (B) dorsal view, and (C) close-up of head of a specimen of Stiliger berghi from Oshoro Bay, Hokkaido, Japan. Arrows indicate characteristic color bands on the rhinophores and the visually distinct esophageal pouch. (D—F) Dorsal and oblique views of Elysia sp. specimens from SW Okinawajima; (G) dorsal view of Elysia sp. from Shirahama, Shimoda, Honshu. Individuals ranged from 3 to 6 mm long. C. D. Trowbridge et al., 2008 Page 51 A. 20 iS 10 5 May 2000 Number of Stiliger berghi Wet Weight (mg) 0 1 2 3 4 5 0 5 10 15 Wet Weight (mg) Length (mm) N on = > © 20 56 ee a 8 oes 2 40 V7 Feel BEE o Ne eriene : BS) [= Polysiphonia o 15 eee tase ; cae = i: 10 te . 20 July 2001 April 2006 2 Z IS =) ZZ 0 1 2 3 4 5 0 5 10 5 Wet Weight (mg) Length (mm) Figure 3. Body size of Stiliger berghi from (A) Sagami Bay in May 2000, (B) Hokkaido in July 2001, and (C, D) Sagami Bay in March 2004 and April 2006. terminal branches, cell size ranged from 200 um wide Polysiphonia sp. (Figure 3B). Specimens on Dasya were and 340 um long to small terminal cells 40 um wide not only more abundant but also significantly smaller and 40 um long. Thus, a wide range of cell sizes was than those on Polysiphonia (Student’s t-test, t = 4.7, available on which slugs could feed. P < 0.001). The overall sacoglossan abundance was In July 2001 at Oshoro Bay, Hokkaido, we found 0.86 slugs per gram (wet weight) of ceramialean algae. high densities of Stiliger berghi on polysiphonous red In feeding-preference experiments, S. berghi preferred algae, primarily Dasya sessilis and secondarily on Dasya to Polysiphonia and strongly preferred both Page 52 The Veliger, Vol. 51, No. 1 Figure 4. (A-C) Radula of a specimen of Stiliger berghi from Oshoro Bay, Hokkaido, Japan. (D) Radula of 2.5 mm Elysia sp. from SW Okinawajima. (E, F) Close-up of radular teeth of two specimens. Scale bars are (B) 50 um, (C) 20 um, (E) 10 pm, and (F) 5 um. C. D. Trowbridge et al., 2008 Dasya and Polysiphonia to Ceramium (Figure 5). All of the initial preferences and two of the three final preferences were statistically significant (Pearson’s chi-square tests, P < 0.05). We observed S. berghi feeding on all these red algae (even Ceramium): many cells were punctured and cytoplasm removed. In March 2004, Stiliger berghi exhibited no prefer- ence between isomorphic female gametophytes (V) and tetrasporophytes (2N) of Polysiphonia harveyana. There was no significant initial or final difference in slug counts between algal life-history stages (Figure 6, Fisher’s exact tests, P = 1.0 for both comparisons). Finally, in April 2008, eight specimens of Stiliger berghi collected from Polysiphonia spp. were observed on the alga Griffithsia japonica in the single-choice trial. We confirmed feeding or attempting to feed in 3 of the 8 specimens. After 2 days, empty algal cells were observed in several containers. Elysia sp. External features: This species was an extremely small plakobranchid with dense white spots on the rhino- phores, dorsal surface of the neck, parapodial edges, and lateral surfaces (Figure 2D—F). The white spots were frequently most dense on the neck and in two patches on the parapodial edge. The red coloration varied among individuals from dark wine-colored to light orange-red, depending on feeding activities (algal host, time since last fed, etc.). The external surface was fairly smooth; there was no evidence of any distinct papillae. In November 2003, we found many individ- uals on red algae with the modal size of 3 mm (Figure 7). The maximum size we recorded at Okina- wajima in seven visits from July 2002 to April 2006 was 6 mm. The single specimen from Shirahama, Shimoda, on the Pacific coast of Honshu, was 8 mm. Although the rhinophores in this one specimen were spaced slightly closer than those in the Okinawan specimens shown (Figure 2), the spacing was within the morphological variation observed in all our specimens. Based on the similarity of the radular formula and highly distinctive tooth shape in the specimens, we consider the Honshu specimen to be conspecific with Okinawan specimens. The radulae of Elysia sp. have 2-3 preradular elements, 10-17 descending teeth (excluding the pre- radular teeth), and 4—5 ascending teeth (including the forming “‘ghost teeth’) for five specimens examined of ca. 1.5 to 2.5 mm in preserved length (see Figure 4D, for example). The leading tooth varied in size: 54 to 69 um in length and 14 to 19 um in basal width. The teeth were blade-shaped with fine denticulation along the cutting edge (Figure 4E—-F). One unusual feature, however, was the blunt tooth tip: it was almost Page 53 rectangular rather than the typical pointed form of most blade-shaped teeth of other sacoglossans. Reproduction and development: The copulatory behav- ior of this species has not yet been observed, but we have recorded spawning in a 6-mm individual. Thus, this Elysia species attained reproductive maturity by at least 6mm in body length. The egg mass was composed of tightly packed capsules with one embryo per capsule. There were small but distinct amounts of white extracapsular yolk distributed within the mass. Because multiple cleavages had already occurred, we were not able to determine ovum and capsule diameters of uncleaved ova. Based on our measure- ments of the existing spawn mass, the embryos were ca. 100 um long and the capsules were ca. 200 um in diameter. Feeding ecology: The ceramialean red algae of Okinawa were extremely diverse, including most of the genera in Table 2; thus, we were not able to document fully all the algal hosts. However, Elysia sp. consumed red algae of several genera, including Griffithsia, Wrangelia, and Polysiphonia. Given the small size of these algae () —-— tetrasporophyte —-©— gametophyte 0 ) 10 15 20 25 Hours Figure 6. Feeding preferences of Stiliger berghi from Sagami Bay, Honshu when offered different algal life-history phases: haploid female gametophytes versus diploid tetrasporophytes of Polysiphonia harveyana. The experiment was conducted in March 2004. Given that many species of Indo-Pacific Elysia species feed either on red algae or both red and green algae, how does our Elysia sp. differ? Elysia furvacauda, E. abeilamakusana, and E. japonica (sensu Jensen) all have black-tipped rhinophores and tail (Table 3); our species does not. Furthermore, the general body coloration of Elysia sp. does not correspond with that of any other superficially similar species, particularly those with orange, red, blue, or yellow spots scattered across the body (Table 3). In terms of internal anatomy (Table 3), Jensen (1985) reported that the radular teeth of E. japonica and E. verrucosa are almost identical (blunt, bladelike teeth) and that Baba (1955) considered E. abei and E. japonica teeth to be similar. However, the blunt, bladelike shape of these species is markedly different from the squared off, almost truncated shape of the specimens of Elysia sp. examined in this study. After investigating all published drawings and SEMs of Elysia radular teeth (worldwide), we consider the teeth of Japanese Elysia sp. to be distinctively different. Furthermore, our species does not correspond in coloration, external morphology, or algal hosts to the insufficiently described Okinawan E. flavipunctata Ichikawa, 1993, or Elysiobranchus ryukyuensis Ichi- kawa, 1993. Not only are our two Japanese red-algal feeders underreported, but most of their ecological analogs around the world are underreported as well (Table 1). Most of these sacoglossans listed are a few mm long, except for Hermaea variopicta (as Hermaeopsis), H. oliviae, H. wrangeliae (both as Aplysiopsis), Elysia abeil amakusana, and E. viridis. The highly branched, delicate ceramialean algae render the red-algal feeders particularly visually cryptic. Many other sacoglossans with dietary homochrony (color derived from algal Page 55 food, resulting in excellent crypsis) are more apparent than the red-algal feeders because of the architecture of the algal hosts. For example, sacoglossans on planar algae (e.g., crustose Codium) or coarsely branched species (e.g., Halimeda, Codium fragile) often are comparatively easy to search for. The darkly pigmented sacoglossans that feed on bright green algae Clado- phora and Chaetomorpha (e.g., see species of Aplysiop- sis Deshayes, 1864, Limapontia Johnston, 1836, and Ercolania Trinchese, 1872) are considerably more apparent than sacoglossans on the uniseriate or polysiphonous filaments of red algae. High densities and local distributions: The abundance and phenology of red-algal feeders is difficult to assess. Hermaea bifida and Stiliger fuscovittatus are considered to be frequent to abundant (e.g., Case, 1972), but most of their ecological analogs are considered to be sparse to rare. If attention were focused specifically on the algal hosts and sacoglossans were expressed on a per hostbasis (number of individuals per gram of host, per algal thallus, or per square meter of algal turf), these cryptic species may be locally quite abundant. Until malacologists standardize abundance values of saco- glossans by the abundance of algal hosts, we will have an incomplete understanding of their occurrence and potential community effects. In Hokkaido, Stiliger berghi was reliably found on ceramialean algae on wave-sheltered shores; prelimi- nary abundance values were 0.86 slugs per gram of host algae (wet weight). In San Francisco Bay, California, S. fuscovittatus occurred on average as 0.48 specimens per ml of Polysiphonia pacifica (wet weight), 2.69 per ml of P. brodiaei, and 0.57 per ml of P. paniculata (Case, 1972). On an area basis, S. fuscovittatus had a maximum local density of 177 specimens per square meter (Case, 1972). Comparable quantitative informa- tion is needed for other red algal feeders. In our study and in that of Case (1972), estimates of population density were determined on a wet-weight basis, not the dry-weight basis used by workers in the Atlantic. Thus, until all the wet-weight to dry-weight ratios have been calculated for all the Japanese hosts, Pacific red-algal feeders cannot be compared with NW Atlantic studies on green algal feeders; however, we do have quantita- tive abundance estimates for >10 Japanese sacoglossan species on green algae (Trowbridge, Hirano, & Hirano, unpublished data). Stiliger berghi and its algal hosts have been recorded in several bays and harbors in close proximity to vectors of introduction (ship hulls and oysters). Its salinity tolerances and those of the ecological equiva- lent species around the world are largely unknown. A notable exception is the NE Pacific Stiliger fuscovitta- tus: Case (1972) reported that the species tolerated salinities ranging from 21 to 33 psu (practical salinity Page 56 The Veliger, Vol. 51, No. 1 # of Elysia on Qo? Wa oe go Whe “yw B. 50 o. 40 “”) © yp S10) = = 20 n=22 SS 40 0 0 1 Z 3.4 52 i Oils: Moe AS Body Length (mm) Figure 7. (A) Specimens of Elysia sp. collected from SW Okinawajima from 2002-2005. (B) Size-frequency distribution of the species in November 2003. Page 57 a a a a ee ee ee a ee sjods poiojoo ApYysiiq ON poddn-yoryq jou [rel poddij-yorfq jou sor10ydouryy Apoq ovs[e s}USUUZTO DSOINAAIA “FT IOAO poio}jeos pure ‘eipodeied jo aspa por snouoydiskjog Ie[npel-sid ¢—-7 pure ‘sauo pue voiuodnl q wWody yUdIOIFIp ‘peoy JO sovjins [esiop ‘so1oydoury.4 oes[e SUIpUsoOSap L[—-O] ‘SuIpusose ¢—p Apounsip ‘yo porenbs opryq jo dry, UO po}eijUI9U09 sjods YSN pol snouoydisouoyy Suoy wim 69-46 4100} SuIpeaT 4199} SYTOpeyq “ung UMOIQ YSIPpoy Apnis sty} Jo ds visdyy_ soroydourys uo sadiiys osiaAsuei} oI L soroydourys 3uriade} ‘suoT Apoq ssoioe sjods Molo ouTyM poop 10 a}1yM eIpodvied jo suisiepy ppaujvoy Peqiiosap JON, Peqliosep 10N ud013 YS] 10 ye pypjoundianyf “7 poddn-yorlq you [rey poddn-yorjq jou soroydouryy s}ods pai pue ‘asueio0 ‘yor[g snose ul deay pur ‘souo pyd4soulojavyy SUIPUSDSOp 6 ‘Y}99} SUIPUdOSe XIS eipodeied puv peoy uo ovyided yunlg psoydopvy]D SUO[ WH OP] 4100) SuIpeaT Y1990} oyTopeyq “unig usois ystAeis Apog DSOINAAIA “FT BINSVIDAIP JO YOR] 0} onp YM so1oydouryl Jo apis [esiop pue peroy jo doy Apoq ssoi9e sJOp on[q pue ‘asuvIO ‘MOTIIA [[eUIS snose UI 499} Jo deoy pure ‘souo suIsivul [eipodeied SUIPUSOSOp [| ‘YI90] SuIpusose xIS UO Uk} IO 914M JO UIO}Ied BezsIZ pydsouojapyy Suo[ wm ¢€/] YI00} SuIpesT Yyq99} dYTPOpe[q “yoous (UMOI ‘Pot ‘Ud9I3) SOTJOIIVA IO[OD JOY L, powuodvl “q DISYIN {IAD Snose UI 499} psoydopvy]g jo deay pure ‘souo Surpusosap Apoq ssoioe sjop pol-asuvIO [TRUS pydsouojavyy €I-IT “YyI901 Surpusose xg [re1 poddn-yor[g sisdodug suo] wm €/] 4300} SuIpeoT 4199] SYIEpe[q s}efNoNUuEd soroydourys poddi-yoelg DUDSNYDUD | 1aGD “7 aes[e poy [re1 poddn-yor[g UOAIIIPOAIIJY oye[nNoNUap soloydourys paddi-yorlg wunipoy PoqLiosap 10N pue poddy iunyjq :Ajoanejua uMOIQ YSIPpoy ppnoopaanf “JO “7 sjsoy [es[e@ UMOUY (9zIs puke S}UNOD) Y}99} Ie[NpeY adeys y100} 1e[npey uoneiojos Apog soroads visdyq ‘Apnys sry} pue (E661) PMLATYO] “(CZ6l) Uasues ‘“(PE61) AoTPuRIg “(SC6I “6r6l) EqrEg Woy ApeuNId paseq sjivjaq ‘saisads asoy) 0} AepIuMIs AT[eOIZO;OYdIow oie IO aeBl[V pol UO pooj VY} sotoeds visd7y oyloe_ UJOIsOM puR o1yIO¥g-opuy Jo uostiedwi0g € 3198.1 C. D. Trowbridge et al., 2008 Page 58 units). The LDs5 9 values (lethal dose for 50% of specimens) for S. fuscovittatus at 10 and 13 psu were 18h and 72h, respectively. Thus, this NE Pacific species could survive in estuaries and bays, predispos- ing it to be inadvertently introduced to other regions (probably accounting for reports from Florida and the Gulf of Mexico). The only other salinity estimate that we have found is 16 psu at the site where Hermaea cruciata was collected in the Atlantic (Vogel, 1971; Marcus, 1972). Reproduction and development: Baba & Hamatani (1952) illustrated and briefly described the spawn mass of Stiliger berghi: the shape (oval), size (max. 4 mm long xX 1mm broad), and coloration (ova were unpigmented). Our results were consistent with their description. Furthermore, our ovum and capsule-size values for S. berghi (Table 1) were consistent with those of Hamatani (1960, 1963). Although not quite as small as those recorded for H. bifida, the ova of S. berghi are among the smaller ones recorded for sacoglossans (Clark & Jensen, 1981; Jensen, 2001). The ovum, capsule, and shell sizes indicate that S. berghi probably has planktotrophic larvae; the type I shell also supports this inference. Our results with the egg mass of Elysia sp. cannot be well compared with the egg masses of other sacoglos- sans, given that we had a single spawn mass and that several embryonic cleavages had already occurred. There were two notable points, however: (1) There was white extracapsular yolk distributed within the mass, comparable with that seen in many other Elysia spp. (Jensen, 2001; Patrick Krug, personal communi- cation). (2) The embryos were relatively small (ca. 100 um), suggestive of planktotrophic larvae (but see Clark & Jensen, 1981; Jensen, 2001). Long-lived planktotrophic larvae (of any inverte- brate) produced within bays, particularly those with commercial shipping, have a greater probability of dispersal via ballast-water uptake and discharge than do short-lived, lecithotrophic larvae or the larvae of open-coast species. Thus, the wave-sheltered habitat, coupled with the long planktonic period of most sacoglossans (typically weeks to months), may predis- pose these species to being introduced accidentally via international trade. The major vector for introduction and movement of ceramialean algae is hull fouling (reviewed by Trowbridge, 2006); thus, sacoglossans that settle on red algae on a ship hull may be unintentionally dispersed within or between ocean basins, depending on the sacoglossans’ tolerance for water currents past the hull and for variations in water temperature and salinity in different embayments. After an initial introduction, species with plankto- trophic larvae would have a greater probability of marginal dispersal via coastal currents or other The Veliger, Vol. 51, No. 1 mechanisms than species with short-lived larvae or with direct development. Thus, red-algal feeders with planktotrophic larvae that dwell in bays may not be as locally endemic as initial reports imply. However, there are internationally supported criteria for recognizing introduced species (reviewed by Trowbridge, 2006) that need to be explicitly considered in evaluating any purported introductions. Feeding ecology: The diet breadth of Japanese red-algal feeders is difficult to evaluate. Baba (1937) recorded the Japanese Stiliger berghi on Zostera weeds (presumably on red-algal epiphytes) and on the red algae Ceramium, Polysiphonia, and Galaxaura (Baba, 1959). However, Baba did not report actual feeding observations but rather field associations, which may or may not reflect trophic associations. Polysiphonia spp. (sensu lato) are frequently sacoglossan hosts, and Ceramium spp. are occasionally hosts. However, Galaxaura is a calcified red alga and probably not consumed by S. berghi. In fact, we have often observed thin strands of green and red filamentous algae among the calcified branches that may well be sacoglossan foods. Thus, we know that S. berghi feeds on at least four of ten genera of ceramialean red algae known to be sacoglossan hosts (Table 2). The subtropical to tropical Elysia sp. has been collected from, and fed on, several algal hosts (this study). The small size (<1 cm) of the intertidal host algae in Okinawajima constrained detailed investiga- tions. Hopefully, future subtidal studies in Japan and SE Asia will elucidate the species’ seasonality and algal- host associations. What is particularly intriguing is that the teeth of Elysia sp. are fundamentally different from those of its ecological counterpart on temperate to boreal shores—namely, Stiliger berghi (Figure 4A— F)—and from all other red-algal feeders worldwide. Sacoglossan species consume red algae on most shores throughout the world. Their feeding habits and diet breadth are extremely difficult to characterize, owing primarily to the taxonomic challenges posed by the algal hosts and secondarily to the insufficientcy of published literature on the subject. The two seemingly well-characterized NE Atlantic sacoglossans exemplify this problem: Hermaea bifida and Elysia viridis (Table 1). Thompson (1976) reported that H. bifida “‘was usually found on red algae, such as_ Griffithsia, Delesseria, and Heterosiphonia’ (p. 174). Garstang (1890) recorded the sacoglossan “creeping over a frond” of Delesseria hypoglossum and stated that the slug would not eat the alga during a 12-day experiment. Yet, Delesseria has persisted as a presumed host in the literature. Other major hosts were not listed—Halurus (e.g., H. equisetifolius) recorded by Duerden (1896) and Bornetia (e.g., B. secundiflora) recorded by Cornet & C. D. Trowbridge et al., 2008 Marche-Marchad (1951) and cited by Miller (1958). Hermaea bifida has been reliably reported from five genera of algae (Table 1), yet the sacoglossan review by Williams & Walker (1999) reported only a single genus (Griffithsia) and the review by Handeler & WaAgele (2007) reported four genera. The literature on Elysia viridis illustrates a compa- rable underestimate. Thompson (1976) reported a number of algae, but many of those listed do not represent algal foods. Typically cited hosts include Codium, Bryopsis, Cladophora, and Chaetomorpha (Williams & Walker, 1999; Handeler & WaAgele, 2007). Yet red-colored E. viridis and the capacity to consume red algae have been known for over a century (Garstang, 1890; Thompson, 1976, and references therein). The sacoglossan feeds on at least four genera of red algae: Griffithsia, Halurus, Dasya, and Dasysi- phonia (Van Bragt, 2004; Trowbridge, personal obser- vations). Consequently, E. viridis feeds on at least eight genera, including green and red algae, native and introduced. With a thorough consideration of the primary literature and enhanced field investigations, we may all improve our estimates of the breadth of the sacoglossan diet. The fact that Hermaea bifida and Elysia viridis have at least five to eight algal genera as hosts is cautionary to proponents of the notion that the breadth of the diet of sacoglossans is narrow. Sacoglossans can and do change algal hosts on spatial and temporal scales (Trowbridge & Todd, 2001; Trowbridge, 2004). Sacoglossans that feed on filamentous algae (green or red) tend to fragment the hosts. With uniseriate green or red filaments (e.g., Cladophora and Chaeto- morpha versus Griffithsia and Halurus), the removal of cytoplasm from a cell can cause structural damage to the alga, resulting in fragmentation. Polysiphonous algae may be slightly more robust, but herbivore- induced damage in wave action contributes to the removal of substantially more algal biomass than the amount consumed (Trowbridge, 1993). Ceramialean algae have a great propensity for their fragments to regenerate and/or reattach. Therefore, sacoglossan herbivory may enhance or exacerbate the spread of introduced red algae in a manner similar to that described for the sacoglossan Lobiger serradifalci (Calcara, 1840) feeding on Caulerpa taxifolia in the Mediterranean Sea (Zuljevic et al., 2001). Kleptoplasty: Of the 20 species of red algal feeders (Table 1), only a few have been evaluated for the functionality of their ingested algal chloroplasts. (1) Hermaea bifida had long-term functional chloroplast retention (Taylor, 1971; Kremer & Schmitz, 1976; Kremer, 1977). (2) The congener H. cruciata exhibited medium-term, nonfunctional retention (Clark et al., Page 59 1990). (3) The survival of starved Stiliger fuscovittatus held in the light versus dark was investigated by Case (1972), but the results were inconclusive. (4) Elysia cf. furvacauda retained functional chloroplasts of both red algae and green algae seasonally (Brandley, 1984) in an intriguing study of seasonal host shift. (5) Finally, there have been extensive studies with Elysia viridis on the species’ long-term, functional retention of chloroplasts of the introduced green alga Codium fragile ssp. fragile (previously called Cf’ ssp. tomentosoides), which have been (reviewed by Williams & Walker, 1999; Trow- bridge & Todd, 2001). However, the capacity to retain chloroplasts of the native green algae Codium vermilara and C. tomentosum (sensu stricto) 1s unclear because the algal hosts were misidentified in many of the classic papers on kleptoplasty of E. viridis; furthermore, the species’ capacity to retain red algal chloroplasts has never been investigated, although the species readily consumed four red algal genera (Table 1). What is the probability that Stiliger berghi and/or Elysia sp. have functional kleptoplasty? Past studies indicate that some of the red-algal hosts have chloroplasts sufficiently robust to maintain functional kleptoplasty. Furthermore, several related sacoglossans have functional kleptoplasty. However, functional kleptoplasty may have evolved multiple times. The assemblage of red-algal feeders needs to be investigated before generalizations can be made. Prospectus: The ecology of sacoglossans that feed either selectively or partially on ceramialean red algae has been understudied, compared with species that feed on green algae. We suggest that there are several priority areas for future research: 1. There is a need to characterize more fully the sacoglossan species that feed on red algae, particularly confirming generic affinity on mor- phological, reproductive, and molecular bases. 2. Future investigators should describe the algal hosts of sacoglossans to genus level (at least) so that diet breadth can be examined more objec- tively. Feeding experiments should be conducted to confirm actual feeding damage and to deter- mine host and nonhost red algae, particularly introduced versus native red-algal hosts. Also, collection and preservation of algal vouchers and their deposition in herbaria would facilitate better future identification of algal hosts. 3. Sacoglossan research is needed at the population level, quantifying the extent of herbivory, based on feeding rates, population abundances, and degree of functional kleptoplasty. 4. The distribution and salinity tolerance of these red-algal feeders need to be documented more fully so the proximity to potential vectors of introduction and the risk of spread (intraocean, Page 60 transocean, and interocean) can be evaluated. With the accelerating frequency of introductions, malacologists should consider whether local spe- cies are indeed native or have arrived from distant shores. Internationally recognized criteria should be used to assess the status of introduced species. Acknowledgments. This study was generously supported by the Sagami Bay Biodiversity Project (2001-2004) by the Misaki Marine Biological Station (MMBS), Women-In-Science Col- laboration (WISC) funds from AAAS/NSF (2002 & 2003), and the Japanese Society for the Promotion of Science (JSPS) Grant-in-Aid for Scientific Research C-15570073 (2003-2006). This paper was written while the first author was supported by the National Science Foundation under Grant No. INT- 0211186. We thank the director M. Morisawa for his extensive logistical support at MMBS; M. Sekimoto and M. Sekifuji for valuable collection assistance in the field; and K. Sayashi for her kind logistical assistance during all our visits. C. Maggs (Queens University, Belfast) kindly identified our Polysiphonia species. We are grateful for the excellent library support by S. Gilmont, J. Mullen, and J. Webster at Oregon State University. We thank D. Behrens, B. Rudman, L. Cervera, P. Krug, C. Carlson, K. Jensen, and H. Wagele for valuable conversations about sacoglossans. Phycological discussions with T. Kitayama (National Science Museum, Tsukuba), M. Yoshizaki (Toho University), W.F. Farnham (University of Portsmouth), and G. Hansen (Oregon State University) substantially improved our understanding of red-algal taxon- omy, particularly as it may relate to sacoglossan feeding. Comments by D. Behrens, K. Brown, and two reviewers substantially improved this paper. LITERATURE CITED ADRIANOV, A. V. & O. G. KUSSAKIN. 1998. A check-list of biota of the Peter the Great Bay, the Sea of Japan. Russian Academy of Sciences, Far East Branch, Institute of Marine Biology, Vladivostock, Dalnauka. BABA, K. 1937. Opisthobranchia of Japan (1). Journal of the Department of Agriculture, Kyushu Imperial University 5:195-235. BABA, K. 1949. Opisthobranchia of Sagami Bay. Iwanami Shoten: Tokyo. 194 pp. BABA, K. 1955. Opisthobranchia of Sagami Bay. Supplement. Iwanami Shoten: Tokyo. 74 pp. BABA, K. 1959. 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GEIGER Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105, USA (e-mail: geiger@vetigastropoda.com) AND TAKASHI OKUTANI Japan Agency for Marine-Earth Science and Technology, 2-15 Natsushima, Yokosuka, Kanagawa Prefecture, Japan 237-0061 Abstract. Anatoma fujikurai sp. nov. is described from the hydrothermal-vent environment at Myojin Knoll, southern Japan. The shell of the species is characterized by the predominant axial sculpture on the shoulder and base and by the undulating selenizone. The animal lacks eyes and shows a radular structure not seen in any other anatomid species examined to date. Hypotheses about its radular structure related to habitat depth, chemosynthetic environment, and geography (Tethys Sea) are rejected; the most plausible explanation is simple interspecific variation. The species is compared with all Japanese anatomid species, as well as to conchologically similar ones, i.e., those with stepped shell profiles from throughout the Indo-Pacific region. Comments on several misidentifications of anatomid species in the literature are provided along with SEM images of the type material of Thieleella sagamiana (Okutani, 1964) and Anatoma soyae (Habe, 1951) for comparison. INTRODUCTION The hydrothermal vent environment (see Van Dover, 2000, for a general review) has yielded many new species, most of which are restricted geographically, as well as in terms of habitat preference, to this unique setting. Many species continue to be described from chemosynthetic environments (e.g., Warén & Bouchet, 2001; Sasaki et al., 2005). The species of small size are less well known in general (for a review of Japanese species, see Sasaki, 2008), which is also the case for deep-sea species. The family Anatomidae contains many undescribed species worldwide (Geiger, 2008). Here, we introduce a new taxon from the Myojin Knoll submarine volcano off southern Japan. The examina- tion of the anatomy of this species, and particularly the radula, has yielded surprising results. MATERIALS AND METHODS Two specimens of the new species were collected from the Myojin Knoll, south off Izu Islands, Japan, 32°06.20'’N, 139°52.17’E, at 1224 m (see Sasaki et al., 2003: fig. 1 for map) on June 24, 2003, on dive HD#185 of ROV Ayper-dolphin during cruise NT03-06 of R/V Natsushima. The anterior parts of the animals were removed from the shells by pulling the head-foot complexes, and the visceral masses remained inside the shells. The isolated animals were photographed under a binocular microscope and dissected to remove the radulae. The shells, opercula, radulae, and animals were examined with a scanning electron microscope (SEM) after being mounted on metal stubs and coated with platinum-palladium. The shells and radulae were cleaned in diluted commercial bleach, and the animals were freeze-dried for SEM examination. Uncoated type specimens of Thieleella sagamiana were examined using variable-pressure SEM. The holotype and paratype of the new species were deposited in the Department of Historical Geology and Paleontology, The University Museum, The University of Tokyo (UMUT). Abbreviations for respositories of specimens and descriptions are as follows: BMNH (The Natural History Museum, London, UK), NSMT (National Museum of Nature and Science, Tokyo, Japan Page 64 [formerly National Science Museum, Tokyo)); SBMNH (Santa Barbara Museum of Natural History, California, USA); SH (shell height); SW (shell width); UMUT (The University Museum, The University of Tokyo, Japan). SYSTEMATICS Class GASTROPODA Clade VETIGASTROPODA Family ANATOMIDAE McLean, 1989 Remarks: See Geiger (2003) for the differentiation of the family from Scissurellidae and Zelaya & Geiger (2007:395) for a diagnosis of the family. The elevation to family rank is based on the molecular phylogeny of Geiger & Thacker (2005, 2006). f Genus Anatoma Woodward, 1859 Type species: Scissurella crispata Fleming, 1828 (orig- inal designation). Anatoma s.1. fujikurai Sasaki, Geiger & Okutani, sp. nov. (Figures 1-4) Type material: Holotype: 3.2 (SW) xX 3.1 (SH) mm (UMUT RM29549), paratype: 3.1 (SW) X 2.7 (SH) mm (UMUT RM29550). Shell: Shell turbiniform, medium size for genus (Figure 1A, C). Protoconch unknown (corroded). Teleoconch I, nine axial ribs on last quarter whorl, no spirals; earlier portion unknown (eroded or broken off). Teleoconch II of 1.75 whorls. Shoulder rounded; approximately 29 comarginal fine, equally spaced axial ribs on first whorl; approximately 10 fine, equally spaced spiral threads running over axial ribs; interstices with fine irregular growth lines. Suture (s, Figure 1A, C) deeply impressed, below selenizone (sz, Figure 1A, C), separated by space equal to width of selenizone. Base rounded, axial ribs (ar, Figure 2A) similar in number and strength on first 1.5 teleoconch I whorls, subsequently becoming less distinct; axial ribs on base slightly more blunt than those on shoulder; approxi- mately 20 fine spiral lines, more or less equally spaced, between selenizone and umbilicus, lines running over axials. Umbilicus (u, Figure 1A) open, deep, moder- ately wide, no funiculus. Aperture rounded, basal adumbilical portion flared (ir, Figure 1A), roof over- hanging. Selenizone slit at periphery (sl, Figure 1A, C), keels (ks, Figure 1C) quite strong, height equal to width of selenizone, undulating, mostly broken off; growth lunules weak; slit open, with parallel margins. The Veliger, Vol. 51, No. 1 Operculum: Operculum (Figure 2B) thin, round, cov- ering aperture, multispiral, nucleus central; margin (om, Figure 2B) is slightly thinner, undulating when dried. Radula: Radula rhipidoglossate (Figure 3), slightly assymmetrical. Rachidian tooth (R, Figure 3B—C, E) taller than wide, with V-shaped cusp, denticle at tip largest, approximately 5 denticles on each _ side, gradually decreasing in size. Lateral teeth 1-3 (L1-3: Figure 3B—C, E) similar, inner edge of cusp smooth, outer edge with 3-4 strong denticles. Lateral tooth 4 (L4: Figure 3E) two-thirds the size of lateral teeth 1-3, cusp triangular, with approximately 10 denticles on inner edge, 4 denticles on outer edge. Lateral tooth 5 (LS: Figure 3C) enlarged by broadening, cusp triangu- lar, denticle at tip largest, approximately 8 on each side, decreasing in size towards base. Inner marginal teeth similar in shape to lateral tooth 5, shaft half the size, cusps decreasing in size towards periphery (Figure 3F). Outer marginal teeth spoon shaped, cusp with many fine denticles. Radular interlock of central field moderate. Animal: Animal’s anterior portion entirely pale in fixed condition except for dark food matter in intestine (i, Figure 2C, D). Head with papillate cephalic tentacles (Figure 4A—-C); papillae all around circumference of tentacle; no eyestalks or eyes (Figure 2C, D); suboptic tentacle (st, Figure 4A, B) short, simple. Epipodial tentacles consisting of four pairs (Figure 4A), tapering toward tips, biserially papillate along anterior and posterior edges (Figure 4E). Epipodial sense organ (ESO, Figure 4A, D) located between second and third epipodial tentacles, stalked, nonpapillte, with blunt tip. Opercular attachment (op, Figure 4A) drop-shaped, with scaly appearance of muscle fibers (Figure 4A, F). Intestine of 1.5 loops visible on surface (Figure 2C, D). Pallial cavity deep; one pair of monopectinate ctenidia attached to pallial roof; hypobranchial gland (hg, Figure 2C) surrounded by loops of intestine in pallial cavity between ctenidia. Anus (a, Figure 2C) opening at anterior central part of pallial cavity. Type locality: Myojin Knoll, south off Izu Islands, Japan, 32°06.20’N, 139°52.17’E, 1224 m. Distribution and type of habitat: Known only from the type locality; hydrothermal-vent field. Etymology: The new specific name is given in honor of Dr. Katsunori Fujikura (Japan Agency for Marine- Earth Science and Technology) for his substantial contribution in biological studies of chemosynthesis- based communities. Remarks: Anatoma fujikurai is distinguished from Japanese species as follows. T. Sasaki et al., 2008 Page 65 Figure 1. Anatoma fujikurai n. sp. Shell. A, Frontal view. Arrowhead indicates end of slit. B, Apical view. C, Back view. D, Basal view. Abbreviations: a = apex; ir = reflected part of inner lip; ks = keels along selenizone; s = suture; sl = slit; sz = selenizone; u = umbilicus. A-B, Holotype, UMUT RM29549. C_D, Paratype, UMUT RM29550. Anatoma lamellata (A. Adams, 1862) from Japan has a similar overall shape and similar density of axial lamellae. However, on teleoconch I, it has a distinct spiral cord in the position of the selenizone (absent in A. fujikurai), teleoconch I is of less then 0.5 whorls (>0.75 in A. fujikurai), the spiral cords are approxi- mately one-third the strength of the axial lamellae (< one-fifth in A. fujikurai), the suture is 1.5 times the width of the selenizone below the selenizone (space equal to width of selenizone in A. fujikurai), and a distinct funiculus is present in the umbilical cavity (absent in A. fujikurai). The differentiation is based on Page 66 The Veliger, Vol. 51, No. 1 Figure 2. Anatoma fujikurai n. sp. A, Enlargement of sulpture of body whorl. B, Exterior view of operculum. C, Dorsal view of anterior part of the animal. D, Lateral view of same part. Abbreviations: a = anus; ar = axial ribs; c = ctenidium; ct = cephalic tentacle; ept = epipodial tentacle; f = foot; hg = hypobranchial gland; i = intestine; mm = mantle margin; om = thin margin of operculum; s = suture; sn = snout; ss = spiral streaks; sz = selenizone. A, Holotype, UMUT RM29549. B-D, Paratype, UMUT RM29550. T. Sasaki et al., 2008 Page 67 Figure 3. Anatoma fujikurai n. sp. Radula. A, D, Six rows of whole radular teeth. B, E, Rachidian (R) and lateral teeth (L1—L4). C, Entire view of rachidian and part of lateral teeth, showing bases of teeth, F, Enlargement of left rows of marginal teeth. A-—C, Holotype, UMUT RM29549. D-F, Paratype, UMUT RM29550. the examination of type material in the BMNH by selenizone, characters not found in A. Jamellata, but SEM (Geiger, pers. obs.). The species has usually been quite typical for T. reticulata; McLean’s (1967: pl. 56, misidentified, e.g., Habe (1951) and Izawa & Matsuoka fig. 8) illustrated specimen is an A. lyra (Berry, 1947); (1999) illustrated an anomphalous species with a Kuroda et al. (1971) showed an unidentified anatomid distinct absence of spiral cords just below the species lacking the lamellae typical for A. /amellata; Page 68 The Veliger, Vol. 51, No. 1 Figure 4. Anatoma fujikurai n. sp. Head-foot of animal. Paratype UMUT RM29550. A, Left lateral view of head-foot with buccal mass removed. B, Enlargement of head. C, Papillate tip of left cephalic tentacle. D, First to second left epipodial tentacles with epipodial sense organ. E, Fourth left epipodial tentacle. F, Surface of opercular attachment. Abbreviations: ct = cephalic tentacles; ESO = epipodial sense organ; LI—L4 = first to fourth left epipodial tentacles; ol = outer lip of mouth; op = opercular lobe; R3-R4 = third and fourth right epipodial tentacles; sn = snout; st = suboptic tentacle. Numanami & Okutani (1990) showed an unidentified species not conspecific with A. lamellata; Okutani & Hasegawa (2000: pl. 18, figs. la, 1b) figured Thieleela reticulata Bandel, 1998. Other published illustrations (e.g., Habe, 1961) are so small as to make positive identification impossible. Only Tsuchida et al. (1991) and Tsuchida & Hori (1996) illustrated the true A. lamellata, and Thiele’s (1912) line drawing agrees exceptionally well with the SEM images of the type material. Anatoma japonica (A. Adams, 1862) from Japan has an overall biconical shape (stepped in A. fujikurai); the density of the axials is at least twice as high as in A. fujikurai, while the spirals are almost as strong as the axials (< one-fifth in A. fujikurai), and the keels of the open slit converge towards the apertural margin (they T. Sasaki et al., 2008 Figure 5. Holotype of Anatoma soyae (Habe, 1951), NSMT Mo-38615. A, Frontal view. B, Basal view. C, Apical view. D, Protoconch. maintain same width in A. fujikurai). The differentia- tion is based on the examination of type material in the BMNH by SEM (D. L. Geiger, personal observation). Anatoma soyae (Habe, 1951) from Japan has an overall biconical shape (stepped in A. fujikurai), has denser axial sculpture (17 vs. 9 on the last quarter of teleoconch I), and the suture inserts immediately below the selenizone on early whorls (space equal to width of selenizone in A. fujikurai), and the keels of the selenizone are low [eroded?] (as high as selenizone width in A. fujikurai). The differentiation is based on SEM images of the holotype (NSMT Mo-38615: Figure 5). Thieleella sagamiana (Okutani, 1964) has a proto- conch with reticulate sculpture, slightly shorter tele- oconch I (0.66 vs. >0.75 whorls) with slightly more axial cords (11 vs. 9 on last quarter whorl). On teleoconch II, the axials are not elevated to low lamellae as in A. fujikurai, and T. sagamiana is anomphalous, while A. fujikurai shows a distinct umbilicus. The comparison is based on SEM examina- tion of the holotype (UMUT RM8808: Figure 6, unfigured in the original description), the paratype (NSMT Mo-69582: Figure 7, Okutani, 1964: pl. 5, fig. 2), and conspecific material collected from southeast off Kamogawa, Chiba Prefecture, 922-959 m (R/V Tansei- Maru, cruise KT-99-6, station 14, 34°48.820’N, 140°37.620'E to 34°48.854’N, 140°39.661’E), Hyuga Basin, 779-803 m (cruise KT-93-9, station HY-4, 31°53'22’N, 131°52'16"E to 31°54'16"N, 131°53'23’E), and Kumano Basin, 2029-2045 m (cruise KT-86-6, Page 70 The Veliger;,, Vol: SIpINowat Figure 6. Holotype of Thieleella sagamiana (Okutani, 1964), UMUT RM-8808. A, Frontal view. B, Basal view. C, Apical view. station KN6, 33°46'6"N 136°40'0"E to 33°46’6’N, 136°37'6"E) (Figure 8). Among Indo-Pacific species with overall stepped shell shape, A. fujikurai is distinguished as follows. Anatoma indonesica Bandel, 1998, has on teleoconch I a spiral cord in the position of the selenizone (absent in A. fujikurai), has more strongly elevated spiral lamellae on the shoulder, crossed by approximately 4-8 fine spiral lines concentrated in the middle of the shoulder (equally distributed in A. fwjikurai), and has fine cancellate sculpture on the base (strong axials and fine spirals in A. fujikurai). Anatoma boucheti Geiger & Sasaki, 2008, from Reunion Island, Indian Ocean, has a teleoconch of 1.125 whorls, shows fine reticulate sculpture of axials and spiral lines of approximately equal strength, and has the keels of the selenizone distinctly elevated. Anatoma herberti Geiger & Sasaki, 2008, from Reunion Island, Indian Ocean, has more strongly elevated axial lamellae and a distinct constriction of the base below the selenizone (absent in A. fujikurai), which bears strongly elevated keels (low/eroded in A. fujikurai) bearing three to four fine axial striae between each axial cord (none in A. fujikurai). Anatoma lamellata nanshaensis Feng, 1996, has on teleoconch I a spiral cord in the position of the selenizone (absent in A. fujikurai), the shoulder is at an angle of approximately 10° (45° in A. fujikurai), the selenizone is in the upper third of the whorl (below the midline in A. fujikurai), and has fine reticulate sculpture T. Sasaki et al., 2008 Figure 7. Paratype of Thieleella sagamiana (Okutani, 1964), NSMT Mo-69582. A, Frontal view. B, Basal view. C, Apical view. D, Protoconch. on the base (more prominent axials in A. fujikurai). The differentiation is based on the original figures of Feng (1996). Anatoma obtusata (Golikov & Gublin, 1978) is overall more depressed (height-to-width ratio = 0.86: 1.03 in A. fujikurai), seems to have reticulate sculpture with axials and spiral of similar strength (axials predominant in A. fujikurai) and has less distinct (eroded?) keels of the selenizone. The differentiation is based on the original line drawing and photographs of the holotype in Kantor & Sysoev (2006). DISCUSSION The generic placement of Anatoma s.l. fujikurai is uncertain, because the diagnostic protoconch sculpture is not preserved in either of the two known specimens. Anatoma s.s. has either smooth or flocculent sculpture, while Thieleella has reticulate sculpture. Whether this character is sufficient to justify generic distinction has been discussed (Marshall, 2002; Geiger, 2003; Geiger & Jansen, 2004; Zelaya & Geiger, 2007) and is provision- ally accepted (see also Geiger, 2006a, b, c). Page 72 The Veliger, Vol. 51, No. 1 Figure 8. Thieleella sagamiana (Okutani, 1964). A-D, Kumano Basin, 2029-2045 m (R/V Tansei-Maru, cruise KT-86-6, station KN6, 33°46'6"N, 136°40'0’E to 33°46'6’N, 136°37'6"E) (SBMNH 83432). E-H, Southeast off Kamogawa, Chiba Prefecture, 922— 959 m (R/V Tansei-Maru, cruise KT-99-6, station 34°48.820'N, 140°37.620’E to 34°48.854'N, 140°39.661’E) (SBMNH 83433). A, E, Frontal view; B, F, Basal view; C, G, Apical view; D, H, Protoconch. T. Sasaki et al., 2008 Figure 9. Radula of Thieleella sagamiana (Okutani, 1964) from specimen shown in Figure 8 E—H. Southeast off Kamogawa, Chiba Prefecture, 922-959 m (R/V Tansei-Maru, cruise KT-99-6, station 34°48.820'N, 140°37.620’E to 34°48.854'N, 140°39.661’E) (SBMNH 83433). A, Central field with inner lateral teeth. Scale bar = 50 um. B—C, Marginal teeth. Scale bar B = 10 um. Scale bar C = 20 um. The shell morphological features clearly place the new species in Anatomidae, including the size, number of whorls for the size, the presence and placement of a slit and selenizone, as well as the common combination of axial and spiral sculptural elements. The presence of strong axial lamellae is unusual in Anatomidae, but it is also known from the Japanese A. lamellata (A. Adams, 1862) and A. herberti Geiger & Sasaki, 2008, from the Indian Ocean. The radula, on the other hand, is exceptional and does not agree with the pattern thus far observed in the family. Although only a handful of species has been examined [A. crispata, A. euglypta (Pelseneer, 1903), A. janetae Geiger, 2006, Anatoma sp. from South Africa, Thieleella argentinae Zelaya & Geiger, 2007, T. baxteri, T. flemingi Marshall, 2002, T. kelseyi (Dall, 1905), T. reticulata Bandel, 1998], in general, they have very similar features also shared with T. sagamiana shown here (Figure 9). These characteristics include a trape- zoidal rachidian tooth with a narrowly V-shaped cusp with the central denticle distinctly larger than the remainder (not broadly V-shaped with denticles of gradually decreasing size); lateral tooth 4 strongly reduced and hook-shaped (not slightly reduced, with distinct cusp bearing multiple denticles); lateral tooth 5 with strongly elongated cusp, at least twice as large as those of the inner marginal teeth, with straight edges and acute angle (not broadened, only slightly larger than the inner marginal teeth, and with convex edges). Although previously it was thought (Sasaki, 1998; Geiger, 2003) that radular morphology will be suitable ‘only for family-level classification—i.e., distinction between Scissurellidae s.s., Anatomidae, Larocheidae, and Temnocinclidae—examination of additional spe- cies in Anatomidae suggests otherwise. Initial indica- tions appeared with A. janetae, whose innermost marginal teeth have very few denticles (Geiger, 2006c) and 4. flexidentata Geiger & Sasaki, 2008, and A. austrolissa Geiger & Sasaki, 2008, which exhibit a radula with floppy teeth. One may hypothesize that the unusual radular form of A. fujikurai is caused by heterochronic process (arrested development). Generally, in other vetigastro- pods, the rachidian and lateral teeth are more narrowly pointed and sharply serrated in juveniles than in adults, as is found in Trochoidea (Warén, 1990) and Halioti- dae (Kawamura et al., 2001), whereas in adults the rachidian often shows an unserrated cutting edge parallel with the row of teeth. Therefore, acutely serrated teeth of A. fujikurai may represent a state of arrested development compared with other typical Anatoma species. This hypothesis, however, should be tested by ontogenetic study in the future. Although we have prepared several hundred scissurellid s.l. radulae covering species over the entire size spectrum (0.6— 11 mm) and including conspecific specimens of differ- ent sizes, we have not seen any ontogenetic differences. Ontogenetic variability is not mentioned in the only modern ontogenetic study of Scissurella by Strasoldo (1991). It is widely acknowledged that food and feeding has a major effect on radular morphology (e.g., Hickman & McLean, 1990). Whether this is the case also in Anatomidae is uncertain as there are no data on food and feeding. Limited gut content analyses have not allowed us to identify specific food sources suggesting deposit feeding (Geiger, personal observation; Sasaki, personal observation). Some alternative explanations for the occurrence of unusual radular morphologies have been explored, but, on the basis of available data, must be rejected. The morphological changes are not correlated with depth, because some deep-water species, e.g., Thieleella argentinae Zelaya & Geiger, 2007, from the abyssal Page 74 plain of the South Atlantic, has the typical anatomid radular pattern. It is tantalizing that two deep-water species from Reunion Island (A. /flexidentata, A. austrolissa: Geiger & Sasaki, 2008) show a very similar yet untypical radular pattern of floppy teeth, but the broader significance is difficult to assess, given the sparse data available. The hydrothermal vent environment is also unlikely as an explanatory factor, as shown by the only moderately modified radula of A. janetae, also collected from sites in the vicinity of hydrothermal vents. The three species with the most aberrant radulae are from the Tethys area: Japan (A. fujikurai) and the Indian Ocean (A. flexidentata, A. austrolissa: Geiger & Sasaki, 2008). The significance of this observation is uncertain because of generally poor sampling of the deep sea owing to a lack of suitable material and the relatively few Atlantic species that have having been examined (Geiger, 2008). The current contribution highlights the wealth of interesting new information that is obtained from small molluscan species. The examination of such understud- ied groups of micromollusks as Anatomidae shows significant promise for the discovery of novel charac- ters and adaptive pathways. Acknowledgments. We thank Dr. Bruce Marshall (Museum of New Zealand Te Papa Tongarewa) and Prof. Carole S. Hickman (University of California, Berkeley) for critically reading the manuscript and helping to improve it. The materials of new species were collected with the kind assistance of the operation team of ROV Hyper-dolphin and the crew of R/V Natsushima (Japan Agency for Marine-Earth Science and Technology: JAMSTEC). Thieleella sagamiana (Figures 8—9) was provided from the late Mr. Eiji Tuschida (formerly of the Ocean Research Institute, The University of Tokyo: ORI) and also collected with support by Prof. Suguru Ohta (formerly of the ORI) and the crew of R/V Tansei-Maru (JAMSTEC, formerly of the ORI). Dr. Kathie Way accommodated DLG during visits to the BMNH. Drs. Tsunemi Kubodera, Hiroshi Saito, and Kazunori Hasegawa assisted us during visits to the National Museum of Nature and Science. This study was supported by a grant-in-aid for Scientific Research from the Japan Society for the Promotion of Science (no. 18340165, 20540455) and by US National Science Foundation grant MRI 0402726. LITERATURE CITED FENG, W. 1996. Microgastropods from Nansha Sea area, China. Pp. 85-205 in Studies on Marine Fauna and Flora and Biogeography of the Nansha Islands and Neighbour- ing Waters II. Ocean Press: Beijing. GEIGER, D. L. 2003. 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ZELAYA, D. G. & D. L. GEIGER. 2007. Species of Scissurellidae and Anatomidae from Sub-Antarctic and Antarctic waters (Gastropoda: Vetigastropoda). Malaco- logia 49:393-443. The Veliger 51(1):76-84 (March 31, 2010) THE VELIGER © CMS, Inc., 2008 Oligocene and Miocene Vesicomyid Bivalves from the Katalla District, Southern Alaska STEFFEN KIEL Institute for Geoscience—Paleontology, Christian-Albrechts-University, Ludewig-Meyn-Str. 10, 24118 Kiel, Germany AND KAZUTAKA AMANO Department of Geosciences, Joetsu University of Education, Joetsu 943-8512, Japan Abstract. Six fossil vesicomyid species from the Katalla district in Alaska are described and illustrated, and three of them are described as new. Calyptogena katallaensis, Archivesica marincovichi, and Archivesica sp. are from the Oligocene Kulthieth Formation, and Archivesica redwoodia and Adulomya spp. A and B are from the lower Miocene Redwood Formation. The Oligocene Calyptogena katallaensis represents the oldest record of Calyptogena, which had previously been traced only into the late Miocene. Archivesica redwoodia shows an unusual mix of characters, including a Calyptogena-like hinge dentition, pallial sinus, and the lack of a nymphal ridge. INTRODUCTION Vesicomyids are a group of highly specialized bivalves that harbor, and rely on, symbiotic sulphophilic bacteria. They thrive in extremely sulfide-rich deep-sea environ- ments, such as hydrothermal vents, methane seeps, and sunken whale carcasses, but they are also found in oxygen-poor and_ sulfide-enriched sediments. Fossil vesicomyids are especially well represented in the uplifted deep-water sediments around the active continental margins of the Pacific Ocean. In a recent evaluation of fossil North Pacific vesicomyids, Amano & Kiel (2007) were able to trace the genus Archivesica back into the middle Eocene (Domengine stage of the California mollusk zones, cf. Prothero, 2001) and to trace Adulomya and Hubertschenckia into the late Eocene. The genus Calyptogena sensu stricto has been traced back into the late Miocene based on literature data (Otatume, 1942; Kanno et al., 1989; Amano & Kanno, 2005; Krylova & Sahling, 2006; Amano & Kiel, 2007). Two fossil vesicomyids have so far been described from Alaska: Archivesica georgemoorei Amano & Kiel, 2007, from the Oligocene? Sitkalidak Formation on Sitkalidak Island (Figure 1), and Adulomya chitanii Kanehara, 1937, from the lower Miocene part of the Yakataga Formation on Kayak Island (Figure 1; Kanno, 1971). The identity of the latter is doubtful and is briefly discussed herein. The purposes of this paper are (1) to describe and to illustrate six species that had been found by others during geologic mapping in the Katalla district of Alaska (Figure 1) in the 1960s and 70s, and (2) to discuss their evolutionary implica- tions. MATERIALS The specimens were collected in the Katalla district by D. J. Miller of the United States Geologic Survey (USGS) at Menlo Park and were reported in a geologic map (Miller, 1975, table 2). Stratigraphically, they are from the Kulthieth and Redwood Formations. The materials are cataloged in the Museum of Paleontolo- gy, University of California, Berkeley (UCMP). The rubber casts of these specimens figured herein were made by F. Stearns MacNeil at the USGS in Menlo Park, California, USA, where these materials were originally housed. Although Miller (1975) assigned preliminary generic identifications to the specimens in his table 2, those identifications were not noted on the specimens themselves. Thus we were unable to correlate Miller’s identifications with our own, except in one case. Despite the fact that the specimens were housed at Menlo Park, the corresponding locality numbers are not Menlo Park numbers (i.e., they do not bear the prefix ““M’’). Kulthieth Formation: Oleinik & Marincovich (2003) summarized the current stratigraphic knowledge of the Kulthieth Formation and suggested that the bulk of the formation is probably early Oligocene in age. The lowermost part of the formation might be Eocene; the uppermost member contains molluscan taxa that are S. Kiel & K. Amano, 2008 ALASKA /'g L Va So ame -gAnchorage ~~~ ie shy 7 Katalla Kayak Island Sess 4 i district Kodiak Isl. en Soo) Sitkalidak Island 442°=W Figure 1. Locality map showing the study area (Katalla district) and the two other places in Alaska from which fossil vesicomyids have been described (Kayak Island and Sitkalidak Island). indicative of the late Oligocene (Oleinik & Marinco- vich, 2003). On Miller’s (1975) geologic map of the central part of the Katalla district, the localities USGS 4312, 4436, 15385, 15387, and 15397 are all plotted together at the site of an abandoned salmon cannery on the north side of the mouth of the Bering River, near the village of Chilkat. The only rock mapped there is ‘“‘Burls Creek Shale/Basin Creek Member undifferenti- ated” of the Kulthieth Formation, and it is considered to be early Oligocene in age. From these localities, we describe Calyptogena katallaensis Kiel & Amano, sp. nov., Archivesica marincovichi Kiel & Amano, sp. nov., and Archivesica sp. Additional specimens from these localities reported by Miller (1975, table 2) include Ancistrolepis teglandae (Weaver, 1942) and Yoldia aff. Y. thraciaeformis (Storer, 1838) at USGS loc. 4312, and Solemya (Acharax) sp. at USGS loc. 4436. Redwood Formation: The Redwood Formation is considered correlative with the lower Yakataga For- mation and thus of early Miocene age (Marincovich, 1990). Three species are described herein from USGS locality 15399 within the Redwood Formation: Archi- vesica redwoodia Kiel & Amano, sp. nov., Adulomya sp. A, and Adulomya? sp. B. Miller (1975, table 2) noted only vesicomyids from this locality. SYSTEMATIC DESCRIPTIONS Family VESICOMYIDAE Dall & Simpson, 1901 Genus Calyptogena Dall, 1891 Type species: Calyptogena pacifica Dall, 1891 (by monotypy); Recent, northeast Pacific. Remarks: Krylova & Sahling (2006) presented a tightly Page 77 defined concept of Calyptogena in which the only fossil species from the North Pacific belonging to Calypto- gena sensu stricto are C. pacifica from northern Japan to the USA, C. panamensis Olsson, 1942, from Costa Rica and Panama, and C. moraiensis Suzuki, 1941, from northern Japan. Among these, C. moraiensis was synonymized with C. pacifica by Otatume (1942); see also Amano & Kiel (2007, table 4). The main characteristics of Calyptogena are a U-shaped cardinal 3 in the right valve, the presence of a nymphal ridge, and the lack of an umbonal pit and of a pallial sinus (Krylova & Sahling, 2006). Calyptogena katallaensis Kiel & Amano, sp. nov. (Figures 2—10) Calyptogena n. sp. 2; Miller, 1975, table 2. Diagnosis: A relatively small, elongate-oval Calypto- gena with short and narrow nymph, well-developed nymphal ridge, and evenly convex ventral margin. Holotype: UCMP 555211, a right valve from USGS loc. 15385, length 19 mm, height 12 mm. Paratype: UCMP 555212, a left valve from USGS loc. 15385, length 45 mm, height 28 mm. Type locality: USGS loc. 15385 in the Katalla district, southern Alaska, USA; Burls Creek Shale Member and organic shale unit, both part of the Kulthieth Formation. Materials: The two type specimens and two additional specimens from the type locality (UCMP 555213, 555214), one specimen from USGS loc. 4436 (UCMP 555217), and two specimens from USGS loc. 15397 (UCMP 555215, 555216), all from the Burls Creek Shale Member and the organic shale unit of the Kulthieth Formation. For measurements, see Table 1. Description: Shell elongate-oval, anterior margin slight- ly more pointed than posterior margin, ventral margin well rounded; sculpture of fine, irregular growth lines. Anterior adductor scar deep, elongate-D-shaped, pallial line distant from shell margin, starts at posteroventral corner of anterior adductor scar, ends at anterorventral margin of posterior adductor scar, no pallial sinus; posterior adductor scar round, weak to moderately deep. Right valve hinge area broad, cardinal tooth | strong, elongate, slightly concave, starting below umbo, subparallel to shell margin; subumbonal cardinal teeth consisting of U-shaped rami (3a and 3b): cardinal 3a thin, elongate, almost parallel to shell margin, with a slight inclination in a anteroventral direction; cardinal 3b broadly triangular with raised edges, anterior raised edge short, length about half of the hinge plate’s height, pointing towards the shell’s interior; posterior edge The Veliger, Vol. 51, No. 1 Figures 2-10. Calyptogena katallaensis sp. nov. from the Oligocene Kulthieth Formation in Alaska. All specimens are rubber peels. Figures 2, 4. Paratype UCMP 555212, left valve, length 45 mm, from USGS loc. 15385. Figures 3, 5: Holotype UCMP 555211, right valve, length 19 mm, from USGS loc. 15385. Figures 6, 8. Left valve from loc. USGS 4436, length 30 mm (UCMP 555217). Figures 7, 9. Right valve, length 29 mm, from USGS loc. 15385 (UCMP 555213). Figure 10. Right valve from USGS loc. 15385, length 28 mm (UCMP 555214). long and straight, area between this edge and nymph moderately deeply excavated. Nymph short and narrow, tapering posteriorly, posterior nymphal ridge present near posterior end of cardinal 3b, but becoming less pronounced during ontogeny. Left valve hinge with moderately thick cardinal 2a, straight and slightly inclined in an anteroventral direction, with thick knob just below umbo; cardinal 2b thin, concave, and pointing in an anteroventral direction; posterior cardinal 4 straight, sharp, almost parallel to shell margin, bordered by a deep, narrow grove below. The largest specimen (paratype UCMP 555212) is 45 mm long and 28 mm high. Discussion: Among the ten Recent species recognized as Calyptogena sensu stricto by Krylova & Sahling (2006) and Krylova & Janssen (2006), the type species C. pacifica is most similar to C. katallaensis n. sp. These two species can easily be distinguished because C. katallaensis has a narrow nymph that tapers posterior- ly, whereas C. pacifica has a broader nymph that has a rather abrupt posterior end. Also similar in general shape is C. valdiviae (Thiele & Jaeckel, 1931), which has a longer nymph than C. katallensis and a much thinner cardinal tooth 3a in the right valve. Distribution: Burls Creek Shale Member and organic shale unit of the Kulthieth Formation, Katalla district, of southern Alaska, USA; upper lower Oligocene. Etymology: Named after the Katalla district in Alaska. Genus Adulomya Kuroda, 1931 Type species: Adulomya uchimuraensis Kuroda, 1931 (by monotypy); Miocene Bessho Formation, central Honshu, Japan. Remarks: We have recently synonymized the elongate vesicomyid genus Ectenagena with Adulomya and raised it to the rank of a genus (Amano & Kiel, S. Kiel & K. Amano, 2008 Page 79 Table 1 Measurements of the specimens. Position of the umbo is % of the total shell length from anterior margin. Minimum values (min.) are given for incomplete specimens. Species Locality Calyptogena katallaensis UCMP 555211 (holotype) UCMP 555212 (paratype) S JSGS loc. 15385 JSGS loc. 15385 e UCMP 555213 USGS loc. 15385 UCMP 555214 USGS loc. 15385 UCMP 555215 USGS loc. 15397 UCMP 555216 USGS loc. 15397 UCMP 555217 USGS loc. 4436 Adulomya sp. A UCMP 555218 UCMP 555219 Adulomya? sp. B UCMP 555220 Archivesica marincovichi UCMP 555221 (holotype) UCMP 555222 (paratype) UCMP 555223 UCMP 555224 UCMP 555225 UCMP 555226 UCMP 555227 UCMP 555228 Archivesica redwoodia UCMP 555229 (holotype) UCMP 555230 (paratype) UCMP 555231 SGS loc. 15399 JSGS loc. 15399 ee ' SGS loc. 15399 SGS loc. 4312 SGS loc. 4312 SGS loc. 4312 SGS loc. 4312 SGS loc. 4312 SGS loc. 4312 JISGS loc. 4312 SGS loc. 4312 CiSieieieieieie SGS loc. 15399 SGS loc. 15399 SGS loc. 15399 UCMP 555232 SGS loc. 15399 UCMP 555233 SGS loc. 15399 UCMP 555234 JISGS loc. 15399 Archivesica? sp. UCMP 555235 SiSieieiats a SGS loc. 4312 2007). The main characters of Adulomya and the hinge features that distinguish it from other vesicomyid genera are shown in Amano & Kiel (2007, fig. 5). Adulomya sp. A (Figures 11-14) Materials: Two incomplete specimens from USGS locality 15399 (UCMP 555218, 555219), Sandstone member of the Redwood Formation, Alaska, USA. For measurements, see Table 1. Description: Shell elongate, beak very far anterior; -anterior muscle adductor scar broadly D-shaped; anterior pedal retractor scar deeply impressed, posi- tioned just anterior of cardinal 2a in left valve and its corresponding socket in the right valve; hinge plate broad, nymph narrow and very elongate, tapering at its posterior end; right valve hinge with two strong Length (L) Height (H) H/L Position of umbo 19 12 0.63 33 45 28 0.62 Dili 29 20 0.69 27 28 20 0.71 33 25 15 0.6 31 31 (min.) 23 — — 30 19 0.63 28 43 20 (min.) — — 53 (min.) 30 (min.) — — 17 (min.) 14 (min.) — — 35 (min.) 20 — — 13 (min.) 10 (min.) — — 31 18 0.58 23 29 (min.) 20 — — 18 (min.) 13 (min.) a 35 20 0.57 24 27 (min.) 17 (min.) = — 29 19 0.65 27 31 17 0.55 28 42 2D 0.52 8 31 (min.) 15 (min.) — — 42 23 0.55 19 31 (min.) 22 (min.) — — 40 19 (min.) — 22 40 19 (min.) — — cardinals, cardinal | nearly vertical, straight, broad, cardinal 3b elongate-triangular, with central ridge: sockets for left valve teeth broad and deeply excavated; left valve with three strong cardinals, cardinal 2a straight, at about 45° to anterodorsal shell margin; cardinal 2b broad, perpendicular to hinge base, fused with 2a below umbo; cardinal 4b elongate, slightly convex; subumbonal pit elongate, deep. Discussion: Because the posterior end is not preserved in any of the specimens, a potential pallial sinus could not be observed. The hinge dentition of Adulomya sp. A is very similar to that described by Kanno (1971, text fig. 10) for Adulomya chitanii Kanehara, 1937, from the Miocene Yakataga Formation on Kayak Island, and the two are likely to represent the same species. However, A. chitanii was originally based on a specimen from Honshu, Japan, and ongoing work on Japanese Adulomya fossils indicates that Kanno’s (1971) Alaskan A. chitanii most probably represents a The Veliger, Vol. 51, No. 1 Tl elelanlelejar=l = = ey Pit Figures 11-16. Adulomya sp. A from the lower Miocene Redwood Formation, southern Alaska, USGS locality 15399. All specimens are rubber peels. Figures 11, 14. Incomplete left valve, 43 mm long (UCMP 555218). Figures 12, 13. Incomplete right valve, 53 mm long (UCMP 555219). Figures 15, 16. Adulomya sp. B from USGS loc. 15399, lower Miocene Redwood Formation, southern Alaska, anterior half of a right valve (UCMP 555220). different species. Unfortunately, Kanno’s (1971) Alas- kan material could not be located at the University of Tsukuba, and the identity of the Alaskan Adulomya species remains uncertain. Adulomya? sp. B (Figures 15, 16) Materials: One incomplete right valve from USGS loc. 15399 (UCMP 555220), Sandstone Member of the Redwood Formation, Alaska, USA. For measure- ments, see Table 1. Description: Only anterior part of shell preserved, but apparently elongate in shape, anterior end parabolic in shape, adductor scar large, broadly drop-shaped, posterior side bordered by a thick, straight ridge; pallial line distant from shell margin, starting just below the center of the posterior side of the anterior adductor muscle scar. Right valve hinge with two strong teeth, cardinal 1 elongate, broad, straight, oriented almost perpendicular to dorsal shell margin with a slight slant towards the posterior; nymph apparently thin and short. Specimen with missing posterior part 17 mm long and 14 mm high. Discussion: This species is currently difficult to place. Its hinge dentition with the reduced cardinal 3a points to a position in Adulomya (compare Amano & Kiel, 2007, figs. 5, 15, 16, 22, and 23), whereas the onset of the pallial line just below the center of the posterior side of the anterior adductor-muscle scar indicates affinities to Archivesica [compare with e.g., A. soyoae (Okutani, 1957), A. kKawamurai (Kuroda, 1943), and A. okutanii (Kojima & Ohta, 1997)]. An extant species that combines both characters analogous with Adulomya? sp. B is ‘Ectenagena’ extenta Krylova & Moskalev, 1996 from Monterey Bay, California. Thus the Alaskan Adulomya? sp. B might be related to ‘Ectenagena’ extenta; the systematic position of both species, however, remains uncertain. Genus Archivesica Dall, 1908 Type species: Callocardia gigas Dall, 1895 (by original designation); Recent, Gulf of California. Remarks: We have recently raised Archivesica to the rank of a genus; its main characters and the hinge features that distinguish it from other vesicomyid genera are shown in Amano & Kiel (2007, fig. 5). Archivesica marincovichi Kiel & Amano, sp. nov. (Figures 17—25) Diagnosis: Elongate-elliptical Archivesica with a thick shell, lunular incision, pallial line starting at poster- oventral margin of anterior adductor muscle scar, and with a small and shallow pallial sinus; cardinal 1 short and thick, cardinal 3a very weak or reduced, cardinal 3b bifid with two short branches. Holotype: UCMP 555221, a right valve of 35 mm length and 20 mm height. Paratypes: UCMP 555222, an incomplete left valve of 13 mm length and 10 mm height. Page 81 S. Kiel & K. Amano, 2008 unular incision Figures 17-25. Archivesica marincovichi sp. nov. from USGS locality 4312, Oligocene Kulthieth Formation, Alaska. Figures 17, 20. Rubber peel of paratype UCMP 555222, left valve, 13 mm long. Figures 18, 19. Rubber peel of holotype UCMP 555221, right valve, 35 mm long. Figure 21, 22. Steinkern and corresponding shell fragment, left valve showing pallial line and adductor muscle scars, 31 mm long (UCMP 555223). Figures 23, 24. Articulated specimen showing sculpture, lunular incision, ligament, and escutcheon, 29 mm long (UCMP 555224). Figure 25. Hinge of right valve (rubber peel, UCMP 555225), note short nymph, section 9 mm long. Type locality: USGS locality 4312 in the Katalla district, southern Alaska, USA; Burls Creek Shale Member and organic shale unit, Kulthieth Formation. Materials: The type material and six additional specimens from the type locality. For measurements and specimen numbers, see Table 1. Description: Shell elongate-elliptical, beak prosogyrate, at anterior third of shell length; lunular incision distinct, broad, occupying three-fourths of anterodor- sal shell margin; ligament about half the length of posterodorsal margin; sculpture of fine, commarginal growth increments. Anterior adductor scar broadly D- shaped, posterior side straight; pallial line somewhat distant from shell margin, with very small and shallow sinus, starting at ventral side of anterior adductor scar, ending at ventral side of posterior adductor scar. Internal ridge from umbo to posterior margin shallow, broadening towards the posterior. Hinge plate strong, right valve hinge with strong but short cardinal | that is oblique to subparallel to the shell margin, cardinal 3a very weak, 3b bifid, anterior branch very thin and short, convex, pointing towards the posterior shell margin, posterior branch slightly thicker and longer than anterior, and also pointing toward the posterior shell margin; nymph relatively short, tapering toward the posterior. Left valve hinge with elongate cardinal 2a that thickens anteriorly, pointing toward the antero- ventral shell margin, cardinal 2b broad, short, pointing and broadening towards the posteroventral shell margin, posterior cardinal 4b strong, elongate, subpar- allel to posterodorsal margin. Discussion: A weak cardinal 3a can be commonly found in A. knapptonensis Amano & Kiel, 2007. Archivesica knapptonensis also has a lunular incision like this new species. However, the surface of A. knapptonensis is sculptured by irregular commarginal growth lines which have not been observed in the new species. In addition, A. knapptonensis has a thinner posterior cardinal 4b in the left valve, and the umbonal pit is better developed than in A. marincovichi. Distribution: Upper lower Oligocene of southern Alaska, USA, known only from the type locality. The Veliger, Vol. 51, No. 1 subumbonal pit Figures 26-31. Archivesica redwoodia sp. nov. from USGS loc. 15399, early Miocene Redwood Formation, Katalla district, Alaska, USA. All specimens are rubber peels. Figures 26, 29. Paratype UCMP 555230, left valve, 31 mm long. Figures 27, 28. Holotype UCMP 555229, right valve, 42 mm long. Figures 30, 31. Paratype UCMP 555231, left valve, 31 mm long. Etymology: After Dr. Louie Marincovich (California Academy of Sciences) who has been studying the molluscan fossils from Alaska. Archivesica redwoodia Kiel & Amano, sp. nov. (Figures 26-31) Diagnosis: Elliptical, moderately elongate Archivesica, right valve with thin cardinal 3a, cardinal 3b bifid, points downward; pallial sinus small, pointed; nymph moderately broad and short. Holotype: UCMP 555229, a right valve from USGS loc. 15399, length 42 mm. Paratypes: UCMP 555230, a left valve from USGS loc. 15399, length 31 mm. Material: The type material and four additional specimens from the type locality. For measurements and specimen numbers, see Table 1. Type locality: USGS loc. 15399, Katalla district, southern Alaska, USA; lower Miocene Redwood Formation. Description: Shell elongate-elliptical, beak at anterior fifth of shell length; anterior adductor scar kidney- shaped in right valve, oval in left valve; pallial line distant from shell margin, starting at posteroventral corner of anterior adductor scar; near posterior margin, it turns upward and ascends towards posterior adductor scar and ends at its anteroventral corner; pallial sinus small and irregular shaped; posterior adductor scar circular except for its almost straight anterior margin; hinge area long and broad, subumbo- nal pit small but deeply excavated in both valves; right valve hinge with strong cardinal 1 that broadens slightly on the lower side and points to the antero- ventral margin of the shell; cardinal 3a thin, elongate, convex, fused with cardinal 3b just below umbo; 3b bifid, anterior branch thin, concave with respect to the anterior shell margin, pointing downwards, posterior branch slightly thicker and longer than anterior branch, straight, parallel to dorsal shell margin; nymph relatively short and moderately broad. Left valve hinge with thick and straight anterior cardinal tooth 2a pointing to the anteroventral shell margin; cardinal 2b very thick, bifid in some specimens, rounded at its posterodorsal side; posterior cardinal 4b long and thin, pointing towards the posterior end of the shell. Discussion: The downward pointing anterior branch of the bifid cardinal 3b in Archivesica redwoodia sp. nov. is unusual for Archivesica. The only other species of Archivesica with such a tooth is A. kawamurai (Kuroda, 1943), but that species differs from A. redwoodia by being larger and by having a slender cardinal 3b and thin cardinals 1 and 2b. The unusual cardinal 3b of A. redwoodia also resembles that of the genera Calypto- gena s.s. and Hubertschenckia. Archivesica redwoodia differs from members of these two genera by the lack of a posterior nymphal ridge, and, in case of Calyptogena, also by the presence of a small pallial sinus. S. Kiel & K. Amano, 2008 subumbonal subumbonal pit WEL Figures 32-35. Archivesica? sp., rubber peels of a steinkern of an articulated specimen (UCMP 555235), length 40 mm, from USGS locality 4312, Oligocene Kulthieth Formation, Alaska. Figures 32, 33. Upper part of left valve. Figures 34, 35. Right valve. Distribution: Lower Miocene Redwood Formation in the Katalla district of Alaska, USA. Etymology: Named after the Redwood Formation in Alaska. Archivesica? sp. (Figures 32-35) Material: One specimen from USGS locality 4312 of the Burls Creek Shale Member and organic shale unit, Kulthieth Formation, Oligocene. For measurements, see Table 1. Description: Shell poorly preserved, very elongate; hinge area broad and short but nymph very elongate and narrow; anterior adductor scar slightly oval, not very deeply impressed, onset of pallial line at its posteroventral corner; hinge of right valve with thin, Page 83 straight cardinal 3a parallel to shell margin; 3b broad, short, pointing to posteroventral corner of shell; cardinal 1 moderately thin, convex with respect to anterodorsal shell margin; left valve hinge with strong cardinals 2a and 2b, cardinal 4b thin, elongate, slightly convex; subumbonal pit well developed just above posterior cardinals in both valves. Discussion: Archivesica sp. differs from A. marincovichi and A. redwoodia described above by its much more elongate shape and nymph. It differs from Archivesica knapptonensis Amano & Kiel, 2007 by having a stronger and longer anterior cardinal 3a in the right valve and a cardinal 3b that is not bifid, and by its thicker anterior cardinal 2a in the left valve. Archivesica georgemoorei Amano & Kiel, 2007, from the Alaskan Oligocene, is not as elongate as Archivesica sp., has a shorter cardinal 3b in its right valve, and a broader nymph. Owing to its poor preservation, we describe this species here only in open nomenclature. DISCUSSION Krylova & Sahling (2006) and Amano & Kiel (2007) traced the fossil history of Calyptogena back into late Miocene time, based on Japanese record of Otatume (1942), Kanno et al. (1989), and Amano & Kanno (2005). Thus, Calyptogena katallaenis sp. nov. from the Kulthieth Formation significantly extends the fossil record of Calyptogena to the late early Oligocene. The rock adhering to the vesicomyid specimens described here consists of siltstone and sandstone; cold- seep carbonate was not seen. Such occurrences outside the hydrocarbon-seep environment received little at- tention in the past; however, they are not rare. Other examples include: Recent Calyptogena sp. from slump deposits on the Laurentian Fan (Mayer et al., 1988); Calyptogena sp. B (= Adulomya) from the upper Pliocene Kurokura Formation in Joetsu, Japan (Amano & Kanno, 2005); Calyptogena pacifica from the Pliocene Kawazume and Nadachi Formations in Joetsu, Japan, and the upper Miocene Morai Forma- tion in Hokkaido, Japan (Kanno et al., 1989; Amano, 2003; Amano & Kanno, 2005); Calyptogena sp. A (= Adulomya) from the middle Miocene Nanbayama Formation, and Adulomya chinookensis (Squires & Goedert, 1991) from a lower Oligocene slump deposits of the Makah Formation in Washington State, USA (Goedert & Squires, 1993). Another potential example is Calyptogena panamensis Olsson, 1942, described from ‘“‘coarse, gritty or pebbly sandstone” (Olsson, 1942, p. 34). Acknowledgments. We thank James L. Goedert (Wauna, Washington) who made us aware of the Alaskan fossils and provided useful comments on the manuscript; Louie J. Marincovich (Californian Academy of Sciences, San Fran- cisco) for his invaluable help with Alaskan Cenozoic Page 84 stratigraphy; and David Haasl (UCMP, Berkeley) for making the material available. Richard L. Squires (CSUN, North- ridge) and an annonymous reviewer are thanked for their helpful reviews. Financial support was provided to SK by a Marie Curie Fellowship of the European Commission (MEIF- CT-2005-515420). LITERATURE CITED AMANO, K. 2003. Predatory gastropod drill holes in upper Miocene cold seep bivalves, Hokkaido, Japan. The Veliger 46:90-96. AMANO, K. & S. KANNO. 2005. Calyptogena (Bivalvia: Vesicomyidae) from Neogene strata in the Joetsu district, Niigata Prefecture, central Japan. Venus 47:202—212. AMANO, K. & S. KIEL. 2007. Fossil vesicomyid bivalves from the North Pacific region. The Veliger 49:270—293. DALL, W. H. 1891. On some new or interesting west American shells obtained from the dredgings of the U.S. Fish Commission steamer Albatross in 1888, and from other sources. U.S. National Museum Proceedings 14:173-191. DALL, W. H. 1895. Diagnoses of new species of mollusks from the west coast of America. Proceedings of the U.S. National Museum of Natural History 18:7—20. DALL, W. H. 1908. The Mollusca and the Brachiopoda. Harvard University, Museum of Comparative Zoology, Bulletin 43:205—487. DALL, W. H. & C. T. Stmpson. 1901. The Mollusca of Porto Rico. United States Fishery Commission, Bulletin 20:351— 524. GOEDERT, J. L. & R. L. SQUIRES. 1993. First Oligocene record of Calyptogena (Bivalvia: Vesicomyidae). The Veliger 36: 72-77. KANEHARA, K. 1937. Miocene shells from the Joban coal field. Bulletin of the Imperial Geological Survey of Japan 27:1-21. KANNO, S. 1971. Tertiary molluscan fauna from the Yakataga District and adjacent areas of southern Alaska. Palaeontological Society of Japan, Special Papers 16:1—154. KANNO, S., K. AMANO & H. BAN. 1989. Calyptogena (Calyptogena) pacifica Dall (Bivalvia) from the Neogene system in the Joetsu district, Niigata prefecture. Transac- tions and Proceedings of the Palaeontological Society of Japan, New Series 153:25-35. Kojima, S. & S. OHTA. 1997. Calyptogena okutanii n. sp., a sibling species of Calyptogena soyae Okutani, 1957 (Bivalvia: Vesicomyidae). Venus 56:189-195. KRYLOVA, E. M. & R. JANSSEN. 2006. Vesicomyidae from Edison Seamount (South West Pacific: Papua New Guinea: New Ireland fore-arc basin). Archiv fiir Mollus- kenkunde 135:231-261. KRYLOVA, E. M. & L. I. MOSKALEV. 1996. Extenagena extenta, a new species of vesicomyid bivalve from Monterey Bay, California. Ruthenica 6:1—10. The Veliger, Vol. 51} 3Nom KRYLOVA, E. M. & H. SAHLING. 2006. Recent bivalve molluscs of the genus Calyptogena (Vesicomyidae). Journal of Molluscan Studies 72:359-395. KurRoDA, T. 1931. Mollusca. Pp. 1-90 in F. Homma (ed.), Shinano Chubu Chishitsu-Shi [Geology of Central Shinano]. Shinano-Kyoiku-Kai: Tokyo. Kuropba, T. 1943. Akebiconcha, a new pelecypod genus. Venus 13:14-18. MARINCOVICH, L. J. 1990. Molluscan evidence for early middle Miocene marine glaciation in southern Alaska. Geological Society of America Bulletin 102:1591—1599. MAYER, L. A., A. N. SHOR, J. HUGHES CLARKE & D. J. W. PIPER. 1988. Dense biological communities at 3850 m on the Laurentian Fan and their relationship to the deposits of the 1929 Grand Banks earthquake. Deep-sea Research 35:1235-1246. MILLER, D. J. 1975. Geologic map and sections of the central part of the Katalla District, Alaska. USGS Miscellaneous field studies map MF-722. OKUTANI, T. 1957. Two new species of bivalves from the deep water in Sagami Bay collected by the RV Soyo-Maru. Bulletin of Tokai Regional Fisheries Research Laboratory 17:27-31. OLEINIK, A. E. & L. J. MARINCOVICH. 2003. Biotic response to the Eocene—Oligocene transition: Gastropod assem- blages in the high-latitude North Pacific. Pp. 36-56 in D. R. Prothero, L. C. Ivany & E. A. Nesbitt (eds.), From Greenhouse to Icehouse: The Marine Eocene—Oligocene Transition. Columbia University Press: New York. OLsson, A. A. 1942. Tertiary and Quaternary fossils from the Burica Peninsula of Panama and Costa Rica. Bulletins of American Paleontology 27:1—106. OTATUME, K. 1942. On the occurrence of fossil Calyptogena from the Ishikari Oil-field, Hokkaido. Journal of the Geological Society of Japan 49:435—437. PROTHERO, D. R. 2001. Chronostratigraphic calibrations of the Pacific Coast Cenozoic: a summary. Pp. 377-394 in D. R. Prothero (ed.), Magnetic Stratigraphy of the Pacific Coast Cenozoic. The Pacific Section SEPM. SQuIREs, R. L. & J. L. GOEDERT. 1991. New Late Eocene mollusks from localized limestone deposits formed by subduction-related methane seeps, southwestern Wash- ington. Journal of Paleontology 65:412-416. STorER, D. H. 1838. Description of a new species of Nucula from Massachusetts Bay. Boston Journal of Natural History 2:122-125. SUZUKI, K. 1941. Three new species of non-marine shells from the Tertiary formations of Hokkaido and Karahuto. Japanese Journal of Geology and Geography 18:53-58. THIELE, J. & S. JAECKEL. 1931. Muscheln der Deutschen Tiefsee-Expedition. II Teil. Deutsche Tiefsee-Expedition 1898-1899 21:159-268. WEAVER, C. E. 1942. Paleontology of the marine Tertiary formations of Oregon and Washington. University of Washington Publications in Geology 5:1—789. THE VELIGER » 9} The Veliger 51(1):85-103 (March 31, 2010) © CMS, Inc., 2008 Trophonella (Gastropoda: Muricidae), a New Genus from Antarctic Waters, with the Description of a New Species M. G. HARASEWY CH Department of Invertebrate Zoology, MRC-163, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013-7012, USA (e-mail: Harasewych@si.edu) GUIDO PASTORINO Museo Argentino de Ciencias Natureles, Av. Angel Gallardo 470, 3° piso, lab. 57, C1405DJR Buenos Aires, Argentina (e-mail: gpastorino@macn.gov.ar) Abstract. The new genus Trophonella is described from the outer shelf and upper continental slope of Antarctica and islands within the Antarctic Convergence. Four previously known species that had been attributed to the genus Trophon (Trophon scotianus Powell, 1951; T. echinolamellatus Powell, 1951; T. enderbyensis Powell, 1958; and T. eversoni Houart, 1997) are included in Trophonella, as is one new species (Trophonella rugosolamellata) described herein. Trophonella resembles Trophon in gross shell morphology: the members of both genera have large, globose shells, paucispiral protoconchs, prominent axial lamellae, and short siphonal canals. Trophonella differs from Trophon in having shells with evenly rounded whorls that lack a well-defined shoulder; rachidian teeth with distinctive, broadly triangular central cusps, but that lack the marginal cusps of Trophon; characteristic spherical accessory salivary glands; and a circumpapillar fold on the penis that is absent in Trophon. Relationships of the genera Trophon and Trophonella, as well as of the subfamily Trophoninae are reexamined by supplementing the data matrix of Kool (1993b, Table 3) with data for additional taxa. Results support the segregation of Trophonella from Trophon at the generic level. Based on the relationships of the type species of their respective nominotypical genera, Trophoninae is either the sister taxon of a narrowly circumscribed Ocenebrinae, or both are part of a larger clade. A better resolved phylogeny containing a much broader sampling of the more than 50 genus-level taxa that have been attributed to these two subfamilies will be required in order to delineate more precisely the membership of the clade and to identify its diagnostic synapomorphies. INTRODUCTION Although the family Muricidae is best known for its temperate and tropical shallow-water representatives, it also includes a group of taxa inhabiting polar seas and intervening deeper waters that have generally been grouped in the subfamily Trophoninae. Cossmann (1903) proposed the subfamily Trophoninae to contain the genera Trophon Montfort, 1810, and Aspella Morch, 1877. The genus Trophon was subdivided into the subgenera Trophon and Trophonopsis Bucquoy and Dautzenberg, 1882. Under Trophon sensu stricto, he included the sections Trophon, Xanthochorus Fischer, 1884, and Forreria Jouseaume, 1880. Trophonopsis was further subdivided into Trophonopsis and Boreotrophon Fischer, 1884. The genus Aspella was not subdivided. Over the past several decades, there has been general agreement among researchers that Trophoninae is not a monophyletic group. As was noted by Radwin and D’Attilio (1976, p. 175), Trophoninae ‘thas been the site of assignment of many dissimilar forms that seem, on the basis of the shell and radular morphology of the type species of Trophon (T. geversianus), unlikely to belong there.” The genus Aspella is presently assigned to the subfamily Muricinae on the basis of radular morphology (Radwin & D’Attilio, 1976, p. 21). These authors went on to exclude from the subfamily Trophoninae a number of genera (e.g., Austrotrophon Dall, 1902; Forreria; Zacatrophon Hertlein & Strong, 1951) because they “... had radulae characteristic of the family Thaididae.” In a phylogenetic study, Kool (1993a, fig. 65) showed that, based on the morphology of its type species, the genus Trophon was closely related to the genera Nucella and Ocenebra, and he suggested that future studies would reveal Trophoninae to be polyphyletic. Our continuing studies on the Muricidae represented in the collections of the United States Antarctic Program (USAP) have revealed that four species previously described in the genus Trophon (i.e., Page 86 Trophon scotianus Powell, 1951; T. echinolamellatus Powell, 1951; T. enderbyensis Powell, 1958; and T. eversoni Houart, 1997) appear to be closely related to each other, but they differ substantially from the type species of the genus Trophon. In this paper, we propose the new genus Trophonella to include these species, and we describe an additional new species that we attribute to this genus. A review of each of the species included in this new genus is provided, based on more recently collected specimens. We also examine the relationships of the genus Trophon and the subfamily Trophoninae by reanalyz- ing Kool’s (1993b, table 3) data matrix of primarily ocenebrine and rapanine taxa supplemented by the addition of data for (1) Trophon geversianus (Pallas, 1769), the type species of the type genus of Trophoni- nae; (2) Ocenebra erinaceus (Linnaeus, 1758), the type species of the type genus of Ocenebrinae; (3) Tropho- nella scotiana (Powell, 1951), the type species of the new genus Trophonella; (4) Boreotrophon aculeatus Watson, 1882; and (5) Paziella pazi (Crosse, 1869), the type species of the genus Paziella. MATERIALS AND METHODS This study is based primarily on specimens collected for the United States Antarctic Program (USAP) aboard the vessels R/V Hero, R/V Eltanin and R/V Professor Sedlacki and housed in the collections of the National Museum of Natural History, Smithsonian Institution (USNM). Additional specimens from several Antarctic expeditions of the Argentine Republic deposited in the Museo Argentino de Ciencias Naturales (MACN) were examined, as were specimens at the Zoological Institute and Museum, Hamburg (ZMH), and the Senckenberg Museum, Frankfurt (SMF), collected by the vessels R/V Walther Herwig and R/V Polarstern. The notation "w/n" following an institutional acronym indicates that no catalog number was associated with the specimen at the time it was examined. All specimens were compared with the holotypes of their respective species, which are housed in The Natural History Museum, London (NHM), the South Australian Museum (SAM), and at USNM. Alcohol-preserved animals of most species of Tropho- nella were dissected, and the gross anatomy of the anterior part of the alimentary system and pallial gonoducts were compared. Radulae were prepared according to the method described by Solem (1972) and examined using a Scanning Electron Microscope (SEM). Terminology for rachidian-tooth morphology follows Kool (1987, fig. 1). Shell ultrastructure data was obtained from fracture surfaces of shell fragments removed from the central portion of the lip along the last shell whorl. In order to determine the generic and subfamilial relationships of this distinctive group of Antarctic The Veliger, Vol. 51, No. 1 muricids, we reanalyzed the data matrix in Kool (1993b, table 3), to which we added data for Tro- phon geversianus, Ocenebra erinaceus, Trophonella scoti- ana, Boreotrophon aculeatus, and Paziella pazi. Harase- wych (1984) was the source of data for T. geversianus, B. aculeatus, and P. pazi. Data for T. geversianus and O. erinaceus were obtained from Kool (1993a). Data on shell ultrastructure of Boreotrophon aculeatus and Paziella pazi were newly obtained from the voucher material from Harasewych’s (1984) study. Each of the newly added taxa were scored for the 18 characters and their character states, as defined in Kool (1993b). The data were analyzed using maximum parsimony (MP) and neighbor-joining algorithms of PAUP* 4.0 Macin- tosh Beta Version 10 (Swofford, 2002). SYSTEMATICS Class GASTROPODA Cuvier, 1797 Order NEOGASTROPODA Wenz, 1938 Family MURICIDAE Rafinesque, 1815 Trophonella new genus Type species: Trophon scotianus Powell, 1951. Diagnosis: Shell large, fusiform, inflated. Protoconch paucispiral (1.5—2 whorls). Teleoconch (6—7 whorls) globose, without distinct shoulder, with tall, conical spire. Sculpture of low, rounded, spiral cords and axial lamellae that may be broadly flaring to short and rugose. Aperture broadly rounded, with flaring outer lip and thin inner lip. Siphonal canal short. Operculum large, oval, with subterminal nucleus. Rachidian teeth with rectangular basal plate; three major cusps, central cusp exceptionally broad, triangular, with small denticle on each side; marginal cusps absent. Accessory salivary glands small, spherical. Esophageal gland reduced. Penis with conical papilla surrounded by collar. Description: Shell large (to 76 mm), fusiform, often broadly ovate, with strongly convex, evenly rounded whorls that do not form a distinct shoulder. Proto- conch paucispiral, occasionally with weak, irregular, spiral cords. Teleoconch sculpture of low, rounded, spiral cords with or without scales, and axial lamellae that range from broadly flaring to short and rugose. Aperture large, suboval, with thick outer lip, and thin inner lip. Siphonal canal short, scabrous. Shell ultrastructure of three layers, the inner two of orthogonally oriented crossed-lamellar aragonite, the outermost layer of calcite. Operculum large, filling aperture, corneous, oval, with subterminal nucleus, thick rim, and large attachment area. Animal large, unpigmented. Cephalic tentacles wide, blunt. Mantle edge smooth, siphon short. Pleuroem- M. G. Harasewych & G. Pastorino, 2008 bolic proboscis short, broad. Radular ribbon small, thin, extending beyond rear of buccal mass. Rachidian teeth with subrectangular, weakly recurved basal plate and smooth, broad, marginal surfaces that lack marginal cusps. Central cusp long, exceptionally broad, triangular; flanked by a single, small to nearly obsolete lateral denticle on each side; lateral cusps short, robust, smooth. Lateral teeth of moderate size, basal plate shorter and narrower than in rachidian teeth, with single scythelike cusp along outer edge that is slightly longer than the basal plate. Esophagus short. Salivary glands very large, ascinous, enveloping esophagus. Accessory salivary glands small, spherical, and com- pletely embedded in corresponding salivary gland. Gland of Leiblein compact, with terminal ampula. Penis large, wide, with conical papilla rising from a concavity at its distal end. Etymology: Trophon (Gr., that which feeds) + ella (L., suffix added to form the diminutive). Included species: Trophonella scotiana (Powell, 1951), new combination [Type species]; 7. echinolamellata (Powell, 1951), new combination; T. eversoni (Houart, 1997), new combination; TJ. enderbyensis (Powell, 1958), new combination; 7. rugosolamellata, new species. Remarks: Trophonella is similar to Trophon in gross shell morphology, as both have globose shells of comparable size with paucispiral protoconchs, promi- nent axial lamellae, and short siphonal canals. Species of Trophonella can be distinguished most readily by their evenly rounded whorls that lack a well-defined shoulder. The main differences appear to be anatom- ical. Trophonella has a circumpapillar fold on the penis that is absent in Trophon. The accessory salivary glands of Trohonella are globular, while those of Trophon are elongate. The rachidian teeth of Trophonella have distinctively broadly triangular central cusps and lack the marginal cusps of Trophon. Trophonella is restricted in geographic distribution to Antarctica and islands within the Antarctic Conver- gence. Records range in depth from 18 m to 474m (both for T. scotiana), with the majority of specimens sampled at outer-shelf and upper continental-slope depths (Figures 20-21). In terms of size and biomass, species of Trophonella are the largest muricids occur- ring within the Antarctic Convergence. Trophonella scotiana (Powell, 1951), new combination (Figures 1-13, 20) Synonymy: Trophon scotianus Powell, 1951, p. 153, pl. 9, figs. 48-49, M88; Carcelles, 1953, p. 189, pl. 2, fig. Page 87 51; Powell, 1960, p. 154; Okutani, 1986, p. 279, pl. 1, figs. 3-6; Dell, 1990, p. 207, figs. 346-347; Castellanos & Landoni, 1993, p. 13, fig. 36; Numanami, 1996, p. 138, figs. 91 a—c. Trophon sp. 1 Hain, 1990, p. 63, pl. 6, fig. 9 a,b; pl. 25, fig. 2. Description: Shell (Figures 1-6) large (to 70 mm), fusiform, thin shelled, chalky. Protoconch (Figures 7— 10) tall, increasing in diameter from 450 um to 1.2 mm in 1'4 to 174 evenly rounded whorls, surface with very fine, irregular and discontinuous spiral threads. Tran- sition to teleoconch marked by broadly flaring lip. Teleoconch of up to 6 slightly globose whorls; spire tall, conical, less than 50% of shell length. Suture impressed to abutting. Spire angle 62°—84°. Aperture subpolygo- nal to evenly rounded, deflected from the coiling axis by 20°—25°; interior glossy. Siphonal canal moderately short, narrow, slightly deflected dorsally. Siphonal fasciole scabrous; pseudoumbilical chink nearly closed or reduced to a very narrow slit. Outer lip rounded with reflected edges. Columellar lip narrow, adpressed. Axial sculpture of regular, thin, strongly developed lamellae (4-11 on last whorl) that span whorl surface from suture to siphon, expanded in shoulder region. Fine, closely spaced growth lines cover whorls and lamellae. Spiral sculpture, of low, rounded, spiral cords, broader than interspaces (18-32 on body whorl, 2-6 on siphonal canal) becoming reduced to obsolete on larger specimens. Shell color white to pale orange- tan. When color present, lighter bands may be evident, one below suture, the other above the juncture to the siphonal canal. Shell composed of three layers (Figure 11). Inner- most layer, of crossed lamellar aragonite with crystal planes oriented perpendicular to the growing edge, comprises roughly 2% of shell thickness. Middle layer, also aragonitic, arranged with crystal planes parallel to growing edge, comprises roughly 36% of shell thick- ness. Outermost layer is calcitic, comprising the major part (62%) of the shell’s thickness. Operculum (Fig- ure 43) oval, with terminal nucleus, outer surface with regular growth lines, inner surface with broad, thickly glazed rim along posterior edge, and large, round/oval attachment area, often with 2-3 horseshoe-shaped scars. Animal large, compact. Foot short, broad, with accessory boring organ situated along ventral midline, opening via ventral pedal gland in females. Tentacles large, broad, closely situated, with large black eyes. Mantle edge smooth, osphradium (Figure 48, os) small, slightly asymmetrical, with 45-55 leaflets per side, less than 50% the length of the ctenidium (Figure 48, ct). Proboscis large, muscular, cylindrical, pleurombolic. Radula (Figures 12-13) short (radular Page 88 The Veliger, Vol. 51, No. 1 Figures 1-11. Trophonella scotiana (Powell, 1951). 1. Apertural, 2. lateral, and 3. dorsal views of a medium-sized specimen with few axial lamellae, USNM 897411, off South Georgia Island, 54°53’S, 35°49’W, in 127-140 m. 4. Apertural view of large specimen with well-developed lamellae, USNM 901644, Off Borchgrevink Coast, Victoria Land, Antarctica, 74°01’S, 178°53’E, in 256— 258 m. 5. Apertural and 6. dorsal views of average specimen, USNM 897560, off South Georgia Island, 54°39’S, 37°22'W, in 140— 150 m. 7-8. Lateral, and 9. apical views of the protoconch. 10. Detail of protoconch sculpture. USNM 901645, W of Graham Land, M. G. Harasewych & G. Pastorino, 2008 length roughly 30% of aperture length), narrow (roughly 550 um wide), extending beyond rear of retracted proboscis. Rachidian tooth subrectangular, basal plate moderately concave anteriorly, with short, smooth marginal area. Central cusp large, broad, lateral cusps shorter, narrower, with single denticle between central and lateral cusps (Figure 13, d) often almost obsolete, but present, even in juvenile speci- mens. Lateral teeth L-shaped, with single cusp along outer edge roughly equal in length to width of basal plate. Juvenile specimens have proportionally larger denticles, and thinner cusps on rachidian and lateral teeth. Esophagus very thin. Valve of Leiblein (Fig- ure 49, vL) ovate. Salivary glands (Figure 49, sg) very large, envelop the valve of Leiblein and the small, nearly spherical accessory salivary glands, joining esophagus via two thick ducts anterior to the valve of Leiblein. Mid-esophagus very thin, lacks conspicuous esophageal gland, passes under gland of Leiblein. Gland of Leiblein large, compact, with blind duct terminating posteriorly in large ampulla. Stomach (Figures 50, 51, sto) large, lining the anterior and dorsal surface of the digestive gland, into which it opens via ducts (Figure 51, dd). Intestine (Figures 50, 51, i) broad. Penis (Figures 45-47, p) large (> 4 X tentacle length), broad, dorsoventrally flattened, elon- gate, with laterally situated sperm duct, dorsal blood sinus, and conical papilla (Figures 46, 47, 52, pp) surrounded by a collar. Pallial oviduct broad. Albumin gland thin, capsule gland (Figure 53, cg) short, globose. Bursa copulatrix (Figure 53, bc) onion-shaped, with distal female opening (Figure 53, fo). Rectum (Fig- ure 53, r) overlays pallial oviduct; rectal gland (Fig- ure 53, rg) lines anterior dorsal surface of rectum; anus (Figure 53, a) adjacent to female opening. Egg capsules of TJ. scotiana lenticular, subcircular in outline, attached to substrate (Hain, 1990, pl. 6, fig. 9b, Trophon sp. 1). Each capsule is about 19mm in diameter, contains 140 eggs that hatch as crawling juveniles in 24-25 months (Hain & Arnaud, 1992, p. 307, table 3). Type locality: Off South Georgia Island, 53°55’S, 38°01'W, in 107 m (Figure 20, CQ). Type material: Holotype, NHM 1961542. Material examined: D indicates empty shells, L indicates live-collected specimens. NHM 1961542, Holotype; USNM 887834, 1 L, off South Georgia Island, 53°50’36’S, 36°18'36"W, 185— 205 m, R/V Islas Orcadas, cr. 575, stn. 30, 19 May oa Page 89 1975; USNM 887835, 1 L, off South Georgia Island, 54°01'18"S, 36°50’42”W, 108-119 m, R/V Islas Orca- das, cr. 575, stn. 24, 17 May 1975; USNM 896099, 4 D, off South Georgia Island, 53°51’S, 37°38’W, in 97— 101 m, R/V Eltanin cr. 22, stn. 1535, 7 Feb 1966; USNM 896982, 1 L, off South Georgia Island, 53°57’S, 36°06’ W, 151-158 m, R/V Professor Siedlecki, cr. 601, stn. 97, 13 Dec 1986; USNM 897411, 2 L, off South Georgia Island, 54°53’S, 35°49’W, in 127-140 m, R/V Professor Siedlecki, cr. 601, stn. 71, 9 Dec 1986; USNM 897416, 1 L, off South Georgia Island, 55°24’S, 35°22'W, 207-218 m, R/V Professor Siedlecki, cr. 601, stn. 60, 8 Dec 1986; USNM 897422, 1 L, off South Georgia Island, 55°08’S, 35°06’W, 145-153 m, R/V Professor Siedlecki, cr. 601, stn. 66, 9 Dec 1986; USNM 897427, 2 L, off South Georgia Island, 55°01’S, 35°27'W, 117-122 m, R/V Professor Siedlecki, cr. 601, stn. 67, 9 Dec 1986; USNM 897433, 2 L, off South Georgia Island, 55°05’S, 35°23’W, 116-121 m, R/V Professor Siedlecki, cr. 601, stn. 68, 9 Dec 1986; USNM 897434, 5 L, off South Georgia Island, 54°31’S, 38°11’W, 180-190 m, R/V Professor Siedlecki, cr. 601, stn. 36, 4 Dec 1986; USNM 897444, 1 L, off South Georgia Island, 53°55’S, 37°08’W, 104-122 m, R/V Professor Siedlecki, cr. 601, stn. 109, 14 Dec 1986; USNM 897459, 1 L, off South Georgia Island, 53°52’S, 37°33'W, 104-112 m, R/V Professor Siedlecki, cr. 601, stn. 111, 15 Dec 1986; USNM 897475, 3 L, off South Georgia Island, 53°52’S, 38°20’W, 113-120 m, R/V Professor Siedlecki, cr. 601, stn. 120, 16 Dec 1986; USNM 897490, 1 L, off South Georgia Island, 54°11’S, 37°53’W, 113-118 m, R/V Professor Siedlecki, cr. 601, stn. 40, 5 Dec 1986; USNM 897521, 2 L, off South Georgia Island, 54°05’S, 38°25’'W, 197-207 m, R/V Professor Siedlecki, cr. 601, stn. 24, 3 Dec 1986; USNM 897560, 1 L, off South Georgia Island, 54°39’S, 37°22'W, 140-150 m, R/V Professor Siedlecki, cr. 601, stn. 46, 6 Dec 1986; USNM 897564, 5 L, off South Georgia Island, 54°51’S, 35°38’W, in 84-103 m, R/V Professor Siedlecki, cr. 601, stn. 72, 9 Dec 1986; USNM 901642, 1 D, Ross Sea, Moubray Pennell Bank, Victoria Land, Antarctica, 73°22’S, 177°37’E, in 465— 474 m, R/V Eltanin, cr. 27, stn. 1933, 30 Jan 1967; USNM 901644, 1 L, Ross Sea, Moubray Pennell Bank, Victoria Land, Antarctica, 74°01’S, 178°53’E, in 256— 258 m, R/V Eltanin, cr. 32, stn. 2018, 14 Jan 1968; USNM 901645, 1 L, W of Graham Land, Palmer Peninsula, Antarctica, 66°21.7'S, 66°47’W, in 70— 106 m, R/V Hero, cr. 731, stn. 1861, 1 Mar 1973; MACN-In 18991, 1 D, Schlieper Bay, South Georgia Island, in 18 m; MACN-In 18990, 1 L, Cumberland Palmer Peninsula, Antarctica, 66°21.7'S, 66°47’W, in 70-106 m. Scale bars = 400 um. 11. Shell ultrastructure. Fracture surface parallel to growing edge. USNM 887835, off South Georgia Island, 54°01'18"S, 36°50’42”W, 108-119 m. Page 90 The Veliger, Vol. 51, No. 1 Figures 12-19. Radulae of species of Trophonella. 12. Dorsal and 13. right lateral views of the radular ribbon of JT. scotiana, specimen in Figure 4. 14. Dorsal and 15. right lateral views of the radular ribbon of 7. echinolamellata, USNM 896041, 2L, NW of Brabant Island, Palmer Archipelago, Antarctic Peninsula, 63°51'S, 62°38’W, in 128-165 m. 16. Dorsal and 17. right lateral views of the radula of 7. enderbyensis, specimen in Figures 38-40. 18. Dorsal and 19. right lateral views of the radula of the holotype of T. rugosolamellata n. sp. ZMH 2777. M. G. Harasewych & G. Pastorino, 2008 Om som [ 100 m 150 m 200m 250m 300 m 350m 400m 450m § 500m be T. scotianus (@) type locality (O) Onna 203 Om 50m 100 m 150m 200 m 250 m 300 m 350 m 400 m T. rugosolamellata (®) T. eversoni (#) type locality & depth unknown Figure 20. Geographic and bathymetric distributions of Trophonella scotiana, T. rugosolamellata n. sp., and the only locality reported for T. eversoni. Bay, South Georgia Island, in 36 m; ZMH w/n, 1 D, off South Georgia Island, 54°13.2’S, 37°49.6’W, in 122 m, R/V Walther Herwig, cr. 68, stn. 15; ZMH w/n, 1 L, off South Georgia Island, 53°55.9’S, 38°29.6’W, in 145 m, R/V Walther Herwig, cr. 68, stn. 19; ZMH w/n, 3 L, 1 D, off South Georgia Island, 53°39.5’S, 37°8.9'W, in 155 m, R/V Walther Herwig, cr. 68, stn. 30; ZMH w/n, 1 D, off South Georgia Island, 53°47.3'S, 37°12.2'’W, in 141 m, R/V Walther Herwig, cr. 68, stn. 31; ZMH w/n, 3 L, off South Georgia Island, 53°59.8’S, 36°56.2’W, in 150 m, R/V Walther Herwig, cr. 68, stn. 35; ZMH w/n, 1 L, off South Georgia Island, 53°43.3’S, 36°26’W, in 230 m, R/V ~ Walther Herwig, cr. 68, stn. 40; ZMH w/n, 1 L, off South Georgia Island, 53°59.1’S, 36°20.6’W, in 198 m, R/V Walther Herwig, cr. 68, stn. 44; ZMH w/n, | L, off South Georgia Island, 53°52.9’S, 36°4.2’W, in 235 m, R/V Walther Herwig, cr. 68, stn. 55; ZMH w/n, 4 L, off South Georgia Island, 54°32.6’S, 35°58.1’W, in 155 m, R/V Walther Herwig, cr. 68, stn. 64; ZMH w/n, 1 L, off South Georgia Island, 54°48’S, 35°23.6’W, in 215 m, R/V Walther Herwig, cr. 68, stn. 70; ZMH w/n, 4 L, off South Georgia Island, 55°15.2'S, 34°46.8’W, in 240 m, R/V Walther Herwig, cr. 68, stn. 78; ZMH w/n, 2 L, off South Georgia Island, 55°0.37’S, 35°03.8’ W, in 124 m, R/V Walther Herwig, cr. 68, stn. 79; SMF w/n, | D, off the Princess Martha Coast, Queen Maud Land, Antarctica, 70°29’S, 8°07'W, in 270-303 m. Literature records: PS ANT V/3 St. 593, 73°55’S, 23°38'W, in 330m (Hain 1990); JARE-25, St. A, 70°14’S, 24°23.9’E, Breid Bay, in 310 m (Okutani 1986; Numanami 1996). Distribution: Off South Georgia Island, the Antarctic Peninsula, and the Ross Sea, the Weddell Sea, and Breid Bay, Antarctica, at depths of 18-474 m. See Figure 20. Page 92 Remarks: This distinctive taxon, the largest species of Trophonella in terms of biomass, had, until recently, been known from less than a dozen specimens (Dell, 1990:208; Numanami, 1996:138). A survey of the collections of Antarctic mollusks at USNM, SMF and ZMH, as well as a search of the literature, has uncovered more than 60 specimens and/or records, most from the vicinity of South Georgia Island and the Weddell Sea. Dell (1990:208) and Numanami (1996:140) both noted variation in shell morphology of this species, with the degree of whorl inflation and the number of axial lamellae per whorl both increased in large specimens. Trophonella echinolamellata (Powell, 1951) new combination (Figures 14, 15, and 21—28) Synonyms: Trophon echinolamellatus Powell, 1951:152, pl. 9, figs. 44, 45. Trophon equinolamellatus [sic] Pastorino, 2002:fig. 22. Description: Shell (Figures 22—24) large (to 69.8 mm), fusiform, inflated, thin shelled, with scabrous surface. Protoconch (Figures 25—27) tall, increasing in diameter from 300 to 920 um in 174 to 2 evenly rounded whorls, surface smooth, frequently pitted. Transition to tele- oconch distinguished by weak flare in lip. Teleoconch of up to 7 evenly rounded whorls; spire tall, conical, slightly less than half the shell length. Suture distinctly impressed. Spire angle 66—72°. Aperture broadly ovate, deflected from the coiling axis by about 25°, interior with glazed, lustrous white layer that does not extend to the end of the flared, rounded, outer lip, nor to the edge of thick, adpressed, columellar lip. Demarcation between apertural glaze and apertural lip very pro- nounced, especially in pigmented shells. Siphonal canal short, narrow, slightly deflected dorsally. Siphonal fasciole scabrous; pseudoumbilical chink pronounced, constricted to varying degrees by thick columellar lip. Outer lip rounded, weakly reflected. Columellar lip thick, overlaying sculpture of previous whorl. Axial sculpture of numerous (14-16 on first teleoconch whorl, 42-50 on last whorl), extremely short lamellae that intersect with closely spaced spiral cords that are as broad as interspaces (16—20 on body whorl, 2-4 on siphonal canal) to produce a scaly surface. On the final whorl of very large specimens, several lamellae may be close together and thickened, giving the appearance of varices that may be spaced '4 to '4 whorl apart. Shell white, or, more commonly pale orange-cinnamon. When color is present, lighter bands may be evident, one below the suture, the other above the juncture to the siphonal canal. The shell is composed of three layers (Figure 28). Innermost layer very thin (approximately 2% of shell The Veliger, Vol. 51, No. 1 thickness), consisting of crossed lamellar aragonite with crystal planes oriented perpendicular to growing edge of the shell. Middle layer of crossed lamellar aragonite, with crystal planes colabrally oriented, accounts for approximately 30% of shell thickness. Outermost layer thickest (~ 68% of shell thickness), calcitic. Operculum (Figure 42) D-shaped, with straight adaxial margin, terminal nucleus, fine growth lines along external surface, and broad, glazed rim along inner surface. Attachment area without scars. The animal is similar to that of T. scotiana in most respects. The radula (Figures 14, 15) is slightly longer (radular length ~ 0.45 aperture length) and has proportionally broader rachidian teeth with more pronounced denticles. Type locality: Off Cape Bowles, Clarence Island, Antarctica, 61°25'S, 53°46’W, in 342 m. (Figure 21, V). Type material: Holotype and one paratype, NHM 1961541. Material examined: D indicates empty shells, L indicates live collected specimens. NHM 1961541, Holotype and one paratype; USNM 638874, 1 L, off Victor Hugo Island, west coast of Palmer Peninsula, Antarctica, 65°08'S, 66°04’W, in 135 m, stn. ED28, 22 Mar 1959; USNM 678397, 3 L, off Palmer Peninsula, Antarctica, 60°48’S, 44°13.5W, in 188 m, R/V Eastwind, stn. EW66-028, 11 Feb 1966; USNM 846180, 1 D, east of Victor Hugo Island, west coast of Palmer Peninsula, Antarctica, 65°04’S, 65°53'’W, in 150m, R/V Polar Duke, 8 Sept 1985; USNM 870316, 1 L, NW of Brabant Island, Palmer Archipelago, Antarctic Peninsula, 63°51'S, 62°38’W, 128-165 m, R/V Eltanin, cr. 6, stn. 439, 9 Jan 1963; USNM 870593, 2 L, NE of Joinville Island, Antarctic Peninsula, 62°41'S, 54°43’W, 210-220 m, R/V Eltanin, cr. 12, stn. 1003, 15 Mar 1964; USNM 881728, | L, off Visokoi Island, South Sandwich Islands, 56°42’18’S, 27°00'24"W, 93-121 m, R/V Islas Orcadas, cr. 575, stn. 61, 30 May 1975; USNM 881906, 1 L, D’Urville Island, Bransfield Strait, Antarctic Peninsula, 62°39’S, 56°10’W, 426-311 m, R/V Eltanin, cr. 6, stn. 418, 2 Jan 1963; USNM 881907, 1 L, D’Urville Island, Bransfield Strait, Antarctic Peninsula, 62°39'S, 56°10’W, 426-311 m, R/V Eltanin, cr. 6, stn. 418, 2 Jan 1963; USNM881939, 1 L 2 D, off South Georgia Island, 54°41’S, 38°38’W, 220-320 m, R/V Eltanin, cr. 9, stn. 671, 23 Aug 1963; USNM 896041, 2 L, NW of Brabant Island, Palmer Archipelago, Antarctic Penin- sula, 63°51'S, 62°38'W, in 128-165 m, R/V Eltanin, cr. 6, stn. 439, 9 Jan 1963; USNM 896056, 2 L, E of South Orkney Islands, 60°51’S, 42°57'W, in 155m, R/V Eltanin, cr. 12, stn. 1083, 14 Apr 1964; USNM 901636, 2 D, off Zavodovski Island, South Sandwich Islands, 56°28.8'S, 27°24.6'W, in 161-210 m, R/V Islas Orca- M. G. Harasewych & G. Pastorino, 2008 100m 150 m 200 m 250 m 300 m 350 m 400m 450 m 500 m T. echinolamellatus (¥) type locality (V) om 50m 100 m 150 m 200 m 250m 300 m 350 m 400 m T. enderbyensis (@) type locality (Q) Figure 21. Geographic and bathymetric distributions of Trophonella echinolamellatus and T. enderbyensis. das, cr. 575, stn. 70, 2 Jun 1975; USNM 901638, 1 L, Gibbs Island, Bransfield Strait, South Shetland Islands, Antarctica, 61°25’S, 56°30’W, in 300 m, R/V Eltanin, cr. 12, stn. 998, 14 Mar 1964; USNM 901639, 1 L, Low Island, South Shetland Islands, Antarctica, 63°26’S, 62°15'W, in 119-124 m, R/V Hero, cr. 691, stn. 26, 10 Feb 1969; USNM 901640, 1 L, Aspland Island, South Shetland Islands, Antarctica, 61°17'S, 56°26’W, in 421-462 m, R/V Polarstern, cr. 2, stn. 22, 21 Nov 1996; USNM 901641, 4 L, Elephant Island, South Shetland Islands, Antarctica, 60°53’S, 55°32’W, in 178-120 m, R/V Polarstern, cr. 2, stn. 79, 9 Dec 1996; col. P. Arnaud, 200-220 m, South Shetland Islands, Antarctica; ZMH w/n, 1 L, NW of Elephant Island, South Shetland Islands, Antarctica, 60°56’S, 55°32.9’'W 81 m, R/V Walther Herwig, cr. 68, stn. 68; ZMH w/n, 2 L, E of Laurie Island, South Orkney Islands, Antarctica, 60°48.9’S, 43°34"W 257 m, R/V Walther Herwig, cr. 68, stn. 87; ZMH w/n, 2 L 2D, E of Laurie Island, South Orkney Islands, Antarctica, 60°41.6'S, 43°57.1’W 290 m, R/V Walther Herwig, cr. 68, stn. 88; ZMH w/n, 1 D, NNE of Laurie Island, South Orkney Islands, Antarctica, 60°34.9’S, 44°17.3'W 240 m, R/V Walther Herwig, cr. 68, stn. 89: ZMH w/n, 4 L, 2 D, SE of Laurie Island, South Orkney Islands, Antarctica, 60°51.3’S, 44°12’W 178 m, R/V Walther Herwig, cr. 68, stn. 90; ZMH w/n, 3 L, 1 D, WNW of Coronation Island, South Orkney Islands, Antarctica, 60°25’S, 46°25.7’'W 150m, R/V Walther Herwig, cr. 68, stn. 118; ZMH w/n, 1L, NW of Aspland Island, South Shetland Islands, Antarctica, 61°21.8'S, 56°0.6’W 368 m, R/V Walther Herwig, cr. 68, stn. 138; ZMH w/n, 1 L, 3 D, NW of Aspland Island, South Shetland Islands, Antarctica, 61°19.8’S, 56°9.9'W 328 m, R/V Walther Herwig, cr. 68, stn. 139; ZMH w/n, 1 L, 1 D, NW of Aspland Island, South Shetland Islands, Antarctica, 61°12.5’S, 56°23.4’W 460 m, R/V Walther Herwig, cr. 68, stn. 141; ZMH Page 94 The Veliger, Vol. 51, No. 1 Figures 22-28. Trophonella echinolamellata (Powell, 1951). 22. Apertural, 23. lateral, and 24. dorsal views of a large specimen, USNM 901636, Off Zavodovski Island, South Sandwich Islands, 56°28.8’S, 27°24.6’W; 161-210 m. 25. Lateral, 26. abapertural, and 27. apical views of the protoconch, USNM 870593, NE of Joinville Island, Antarctic Peninsula, 62°41'S, 54°43’W, 210-220 m. 28. Shell ultrastructure. Fracture surface parallel to growing edge. USNM 901638, off Gibbs Island, Bransfield Strait, South Shetland Islands, Antarctica, 61°25’S, 56°30’W, in 300 m. M. G. Harasewych & G. Pastorino, 2008 w/n, 2L, W of Elephant Island, South Shetland Islands, Antarctica, 61°12.7'S, 55°56.4’W 134 m, R/V Walther Herwig, cr. 68, stn. 148; ZMH w/n, 2 L, W of Elephant Island, South Shetland Islands, Antarctica, 61°8’S, 55°56.2’W 125 m, R/V Walther Herwig, cr. 68, stn. 150; ZMH w/n, 1 L 1 D, NW of Elephant Island, South Shetland Islands, Antarctica, 60°53.9’S, 55°30.4’W 135 m, R/V Walther Herwig, cr. 68, stn. 159. Literature records: Only the type locality has been reported in the literature previously. Distribution: Known from Antarctic Peninsula, South Shetland Islands, South Georgia Island, South Sand- wich Islands and South Orkneys Islands, at depths ranging from 93 m to 460 m (Figure 21). Remarks: The characteristic sculpture of the shell, consisting of sharply raised spiral cords overlain by closely spaced lamellae to produce a scabrous surface, distinguish- es T. echinolamellata from all other known species of Trophonella. Trophonella echinolamellata and T. scotiana have similar bathymetric ranges, and they overlap in portions of their geographical ranges. However, T. echinolamellata appears to be restricted to the Scotia tectonic plate, based on the material thus far available. Trophonella enderbyensis (Powell, 1958), new combination (Figures 21, 29-31) Synonymy: Trophon enderbyensis Powell, 1958:197, pl. 3, fig. 1. Description: Shell (Figures 29-31) large (to 70 mm), fusiform, moderately thin-shelled, with regular spiral sculpture and widely spaced lamellose varices. Proto- conch and transition to teleoconch unknown. Tele- oconch with 7 or more evenly convex whorls; sutural shelf obsolete; spire tall, narrow, conical, approximate- ly 50% of shell length; suture abutting; spire angle about 55°. Aperture broadly ovate, deflected from the coiling axis by 25°, interior glossy white. Siphonal canal short; siphonal fasciole conspicuous, with recurved, scabrous processes; pseudoumbilical chink reduced to a narrow slit. Outer lip rounded, reflected; columellar lip thin. Axial sculpture of low, generally distantly spaced thin lamellae covering entire whorl, obsolete on early whorls, approximately 3 per whorl except for the final whorl, where 3 lamellae are closely adjacent in the ~ holotype. Spiral sculpture of low, rounded cords (3 or 4 on early whorls, 20 on last whorl), about as wide as interspaces, which may have 1-4 fine spiral threads between adjacent cords. Shell whitish. Shell composed of three layers, as in T. scotiana. Operculum (Fig- ure 44) oval, with terminal nucleus; outer surface with Page 95 regular growth lines, inner surface with broad, thickly glazed rim along posterior edge. A single, male animal was examined and found to be anatomically similar to TJ. scotiana. The radula (Figures 16-17) is similar to that of T. scotiana but has more prominent denticles between the central and lateral cusps of the rachidian. Type locality: (Figure 21:7), off Enderby Land, Antarctica, 65°48’S, 71°24’E in 193 m. [British, Aus- tralian, and New Zealand Antarctic Research Expedi- tion, St. 41, January 24/25 1930]. Type material: Holotype, SAM D15497. Material examined: Holotype; SMF w/n, 1 D, Off the Princess Martha Coast, Queen Maud Land, Antarc- tica, 70°30’S, 8°4’W, 261—263 m; SMF w/n, 1 L, off the Princess Martha Coast, Queen Maud Land, Antarc- tica,71°23’S, 13°58’W, 293-357 m. Literature records: None, apart from the original description. Distribution: Presently known from three specimens, one from type locality (off Enderby Land) and two from off Queen Maud Land, Antarctica. Remarks: In his description of Trophon enderbyensis, Powell (1958:198) noted that this species resembled T. scotianus, which, at the time, was known only from South Georgia Island. The specimens presently avail- able indicate that the geographic and bathymetric ranges of these two species overlap. Both Dell (1990:208) and Numanami (1996:140) commented that larger specimens of T. scotiana tend to have an increased number of axial lamellae and a more inflated final whorl of the shell. The adult holotype of Trophonella enderbyensis is comparable in size to the largest specimens of JT. scotiana, yet it has far fewer, shorter, axial lamellae and a narrower, more fusiform shell. The intermediate denticle on the rachidian teeth of T. endebryensis is pronounced, but it is reduced or absent in JT. scotiana. As more material becomes available, especially from off Enderby Land and Wilkes Land, the status of T. enderbyensis may require reevaluation. Trophonella eversoni (Houart, 1997) new combination (Figures 32—34) Synonymy: Trophon eversoni Houart, 1997:9, figs. 1—2, 4, 6. Description: Shell (Figures 32-34) large (to 75.8 mm), fusiform, of medium thickness, strongly sculptured with spiral cords and axial lamellae. Protoconch and transition to teleoconch unknown. Teleoconch with 6 Page 96 The Veliger, Vol. 51, No. 1 Figures 29-37. Trophonella enderbyensis (Powell, 1958). 29. Apertural, 30. lateral, and 31. dorsal views of the holotype, SAM D15497, off Enderby Land, Antarctica, 65°48’S, 71°24’E in 193 m. 32-34. Trophonella eversoni (Houart, 1997). 32. Apertural, 33. lateral, and 34. dorsal views of the holotype USNM 880177, South Atlantic Ocean, ? Antarctica, ? Kerguelen Is. 35—37. Trophonella rugosolamellata new species. 35. Apertural, 36. lateral, and 37. dorsal views of the holotype, ZMH 2777, off South Orkney Islands, 60°52.2'S, 44°28.4'W, in 205 m. Page 97 M. G. Harasewych & G. Pastorino, 2008 Figures 38-46. Trophonella enderbyensis (Powell, 1958). 38. Apertural, 39. lateral, and 40. dorsal views of a young specimen, SMF w/n, off the Princess Martha Coast, Queen Maud Land, Antarctica, 71°23'S, 13°58’W, in 293-357 m. 41. T. rugosolamellata new species, operculum of holotype, Figures 35—37. 42. T. echinolamellata (Powell, 1951), operculum of specimen in Figures 22—24. 43. T. scotiana (Powell, 1951), operculum of specimen in Figure 4. 44. 7. enderbyensis (Powell, 1958) operculum of specimen in Figures 38-40. 45-46. Critical-point dried penis of T. scotiana (Powell, 1951). 45. Dorsal and left lateral views of penis. 46. Detail of terminal papilla. Page 98 The Veliger, Vol. 51, No. 1 90 Figures 47-53. Trophonella scotiana (Powell, 1951). 47. Anterior portion of male specimen, mantle reflected. 48. Ctenidium and osphradium. 49. Anterior alimentary system. 50. Dorsal and 51. right lateral views of stomach. 52. Distal tip of penis. 53. Pallial oviduct. Abbreviations for these figures: a, anus; asg, accessory salivary gland; bc, bursa copulatrix; cg, capsule gland; ct, ctenidium; dd, ducts to digestive diverticula; e, esophagus; fo, female opening; i, intestine; mo, mouth; os, osphradium; p, penis; pp, penial papilla; r, rectum; rg, rectal gland; s, siphon; sg, salivary gland; sto, stomach; vl = valve of Leiblein. M. G. Harasewych & G. Pastorino, 2008 or more globose whorls. Sutural shelf present in last whorls. Spire conical, about 33% of total shell length. Suture abutting. Spire angle about 56°—59°. Aperture large, suboval, deflected from the coiling axis by 27°: interior glossy white; siphonal canal moderately short, slightly inclined; siphonal fasciole scaly. Outer lip rounded with reflected edges; columellar lip thin, tightly adpressed, pseudoumbilical chink absent. Axial sculpture of pronounced, regularly spaced, thin lamel- lae, 12-14 on last whorl, running along whorl surface from adapical suture to the siphonal fasciole. Spiral sculpture of low, rounded, spiral cords: 3, 8, and 24 on the first, second, and last whorls, respectively. Cords most pronounced along shell periphery, reduced near suture and siphonal canal. Cords cover the lamellae but became obsolete along their edges. Shell opaque white. Regular growth lines covering whorls and lamellae. Operculum, radula, and anatomy unknown. Type locality: (Figure 20:¢), Antarctica, 185 m. Material examined: Holotype, USNM 880177. Literature records: Kerguelen Islands (Paratype 2): submarine bank “‘South” off the Kerguelen Islands, in “approximately” 250 m (Houart, 1997). Distribution: This species is based on a holotype collected by a Russian trawler “in the Antarctic,” from a depth of 185 m. Paratype 2 is reported, with some certainty, to have been taken off the Kerguelen Islands. Houart (1997:9) mentions two additional specimens from a submarine bank south of the Kerguelen Islands, at depths of 250 m. Remarks: Trophonella eversoni is known from five specimens, four of which are in private collections. The geographic and bathymetric distributions of this species are apocryphal, and they have yet to be confirmed by subsequent collections. Powell (1957:113) reported the occurrence of five species of Trophon in the Kerguelen Islands, although all but two of the stations sampled by the British, Australian, and New Zealand Antarctic Research Expedition were from substantially shallower depths. Houart (1997:11) noted that T. albolabratus Smith, 1875, one of the Kerguelen Island species, differs from T. eversoni in having “low, thin, and more numerous axial lamellae, stronger spiral cords, and a shorter siphonal canal. He distinguished T. eversoni from T. scotiana on the basis of the former having more numerous spiral cords, more numerous and abapertu- rally sloping axial lamellae that are not as strongly produced adapically. As with T. enderbyensis, addi- tional collections from off Enderby Land, Wilkes Land, and the islands off their coasts will be required to assess more accurately the distribution and relation- ships of T. eversoni. Page 99 Trophonella rugosolamellata new species (Figures 20, 35—37) Description: Shell (Figures 35-37) large (to 76 mm), fusiform, solid, chalky, with strong spiral cords and thick axial lamellae that are furrowed and reflected at the shoulder. Protoconch unknown. Teleoconch of at least six narrowly convex whorls; spire 33% of total shell length. Suture abutting. Spire angle about 59°. Aperture large, oval, deflected from the coiling axis by about 22°, interior glossy white. Siphonal canal moderately long and narrow, slightly inclined, siphonal fasciole strongly scabrous, pseudoumbilical chink closed. Outer lip rounded with reflected edge. Colu- mellar lip broad, with thick, expanded callus. Axial lamellae thick, more or less evenly spaced (11 on last whorl, 8 on the penultimate whorl), running along whorl surface from suture to the siphonal fasciole, narrowest at midwhorl, most pronounced near and on the siphon and especially at a position corresponding to the shoulder, where they are enlarged, furrowed and strongly reflected. Regular growth lines evident on the whorls and lamellae. Spiral sculpture of rounded, closely spaced cords (3 or 4 on early whorls, 20 on last whorl) that extend onto the outer surfaces of the lamellae. Cords most pronounced from siphonal canal to region corresponding to the shoulder. Shell color whitish. Shell composed of three layers, as in T. scotiana. Operculum (Figure 41) subpolygonal, thick, brownish, with terminal nucleus. Growth lines on external surface; broad, glazed rim covering nucleus; attachment area smaller. Animal large. Foot short, broadly rectangular. Tentacles very wide, blunt, with small black eyes; mantle edge smooth, siphon very short. Osphradium small, asymmetrical, about 50% ctenidium length. Pleurembolic proboscis short and broad. Radular ribbon (Figures 18, 19) short (radular length ~ 0.44 aperture length), thin, extending beyond rear of buccal mass. Rachidian tooth subrectangular, and basal plate weakly concave anteriorly. Central cusp broadly triangular, flanked by short stout denticles. Outer cusps shorter and narrower, slightly outwardly direct- ed. Marginal cusps absent, marginal area smooth. Lateral teeth L-shaped, narrower than rachidian teeth. Esophagus short. Salivary glands large, ascinous, whitish, with ducts entering esophagus just anterior to the wide valve of Leiblein. Accessory salivary glands small, spherical, located in the anterior part of the buccal mass, totally embedded in salivary glands. Gland of Leiblein compact, without lobules, brownish in color, overlaying esophagus. Penis similar to that of T. scotiana; large, wide, flat, > 4 times tentacle length; with terminal, conical papilla surrounded by narrow collar. Page 100 Muricanthus Forreria Nucella Acanthina Trochia Ocenebra erinacea Trophon geversianus Haustrum The Veliger, Vol. 51, No. 1 Ocenebrinae Trophonella scotiana Boreotrophon aculeatus Paziella pazi Morula Cronia Vexilla Drupa Nassa Pinaxia Dicathais Vasula Tribulus Thais Neorapana Mancinella Purpura Plicopurpura Rapana Stramonita Concholepas MP Cymia Rapaninae NJ Figure 54. Strict consensus maximum parsimony [MP] and neighbor-joining [NJ] trees depicting the relationships of genera based on data derived from their type species. Data from Kool (1993b:Table 3) have been supplemented by data for Ocenebra erinaceus, Trophon geversianus, Trophonella scotiana, Boreotrophon aculeatus, and Paziella pazi. Type locality: S of Laurie Island, South Orkney Islands, 60°52.2'S, 44°28.4'W, in 205 m. Type material: Holotype, ZMH 2777. Material examined: This species is presently known only from the holotype. Distribution: Known only from the type locality, collected by the R/V Walther Herwig cr. 68, stn. 91 (Figure 20). Etymology: rugosolamellata from Latin rugosus and Latin lamellata (with lamellae, or thin plates) referes to the extension of the spiral cords onto the axial lamellae. Remarks: Trophonella rugosolamellata most closely resembles 7. eversoni, but it can be distinguished by its thicker shell, with slightly fewer lamellae and fewer, coarser spiral cords. The axial lamellae of T. rugoso- lamellata are distinctive in that they are reflected along the outer edges, especially at a position corresponding to the shoulder, where they are enlarged, furrowed, and M. G. Harasewych & G. Pastorino, 2008 Page 101 Table 1 Taxa, characters, and character states used in an analysis of the relationships of Trophoninae. The data matrix is that of Kool (1993b:table 3), to which have been added the taxa Trophon geversianus, Ocenebra erinaceus, Trophonella scotiana, Boreotrophon aculeatus, and Paziella pazi. See Kool (1993b) for detailed descriptions of characters and character states. N Ww aN n Character 1 6 Taxa from Kool (1993b): Muricanthus Forreria Nucella Haustrum Morula Cronia Rapana Cymia Stramonita Concholepas Dicathais Vasula Vexilla Nassa Pinaxia Drupa Plicopurpura Thais Purpura Mancinella Neorapana Tribulus Acanthina Trochia ToutrvvVvvVv VA vVewOnmnarvdarsreavrearsrvoggyn Tr rQ QnOaanawrwyvreaevnanawoerrqgtmoonrrnraoecges Toca cMWMAAAAHAAqgDqAqgAAAATrTA TH wVvVROOHOHMPAHAHAPRRRAHRARAAA GAH &© Oo w SCOP PDP VOOR VHA HHP VOPR WHO MAVOETHD PegoogoCP DM DP PS THM OP P OY OY PL OH O P PL Dw Newly added taxa: Ocenebra erinaceus Trophon geversianus Trophonella scotiana Boreotrophon aculeatus Paziella pazi crAac;ge eeorne qaaqnavoetes ooo & Pp oxo oP & O35) OS) 6) O36)-ORS) strongly reflected to produce a tapering structure similar to the open shoulder “spine” of Pterochelus Jousseaume, 1880. The spiral cords of T. eversoni are more numerous and thinner, and they are not as pronounced along the siphonal canal or on the axial lamellae as in T. rugosolamellata. Trophonella ender- byensis has a higher spire, fewer, smaller, and more widely spaced lamellae, and fainter spiral cords than T. rugosolamellata. PHYLOGENETIC RELATIONSHIPS A maximum parsimony analysis of the data matrix shown in Table 1 [PAUP* 4.0 beta version 10 Heuristic search, Accelerated transformation, TBR Swapping algorithm] produced 475,972 equally parsimonious trees (length = 60; consistency index = 0.783; retention index = 0.903]. A strict consensus of these trees is 7 yor nrnennnnnnnvnvanvwevreereeaegrerrpere FOOD eaoon oe 8 \o = So = to Ww = i WN = on = NX 0 fe) yrvivorTroooooocoootoomtrprperpe tee P & & Ot © © © © © 2 © © Wl © Wl © © © © © FL © WL © © © WV a ed VVVAaNQ HAMM AACAACHAMAHOPAgTTHR Ho pw wyvvavaanananaawvaawvnandngggn YoFOOTR O97 AAA M4 HOAACAARAAATTD PD wyeveaananaanananananhananknagoa sa ®& yMvvVvOortooovTooooToooo oO oO ke fm Pm OD yVvVVAOHnNAAnAnAnnAanacagctegcatcgcaoagcge VVVP PPP HPHHP HH VO PHPAPHRA ATTY PVP HPHPHHMPHHPHPHPHPHPHHPHPHRPDH OOD VV oe FOr Se ee AHO TOO ff » © P p 2 © P © p © © P eoooes Sooo s pppoe p © P eooa0c Tooose peeaoge »Pppoad shown in Figure 54 (MP). Figure 54 (NJ) shows the neighbor joining tree based on the same data. Ocenebra erinaceus, the type species of the type genus of Ocenebrinae, emerges in an unresolved pentatomy with the genera WNucella, Trochia, Forreria, and Acanthina in the strict-consensus tree and as sister taxon to these genera in the neighbor-joining tree. Trophon geversianus, the type species of the type genus Trophoninae, emerges as the sister group to this clade in both the MP analysis and the NJ analysis. While the NJ tree shows Haustrum to be the sister taxon to this clade, which, in turn, was joined by the clade ((Trophonella + Boreotrophon) FPaziellaj, the MP analysis groups the genera Haustrum, Trophonella, Boreotrophon, and Paziella into an unresolved polyt- omy that is sister to Trophon and the polytomy containing Ocenebra. These results indicate that the type species of Page 102 Trophonella is more closely related to Boreotrophon than to the type species of Trophon and thus sup- port the segregation of Trophonella from Trophon at the generic level. They also confirm the conten- tions by various authors that Trophoninae, as defined by Cossmann (1903:9), is not monophyletic. Based on the relationships of the type species of their respective type genera, either Trophoninae is the sister taxon of Ocenebrinae or both are part of a larger clade. In his phylogenetic study, Kool (1993b:fig. 30) considered Ocenebrinae to include Haustrum. Tan (2003) subsequently erected the subfamily Haustrinae (with Haustrum as the type genus) to encompass a clade of nonrapanine muricids endemic to Australia and New Zealand. However, the relationship of this clade to Ocenebrinae and Rapaninae varied significantly, depending on the choice of outgroup. Beu (2004:216) concluded that it seemed preferable to retain the taxa included by Tan in Haustrinae within the subfamily Ocenebrinae. This broader delineation of Ocenebrinae includes the type species of Trophoninae in both MP and NJ trees. The genera Trophonella and Boreotrophon emerged as part of an unresolved polytomy with Haustrum in the strict-consensus MP tree, but were depicted as being more distantly related in the NJ tree. Both Ocenebrinae and Trophoninae were proposed in the same publication (Cossmann, 1903:10), so neither has priority. Although it is likely that these two subfamilies will be synonymized, based on the close relationship between the type species of their type genera, a better resolved phylogeny, including a more comprehensive sampling of the more than 20 genus- level taxa attributed to each of these two subfamilies, will be required to delineate more precisely the membership of the clade and to identify its diagnostic synapomorphies. Acknowledgments. We thank the following curators and collection managers for access to specimens in their collec- tions: K. Way (NHM); A. Warén (NHRM); P. Bouchet and V. Heros (MNHN); P. Mikelsen (AMNH); B. Hausdorf (ZMH), A. Tablado (MACN); C. Ituarte; (MLP) and R. Janssen (SM); P. Arnaud, Endoume, France. This work was supported by a grant from the Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET), Argen- tina, which enabled the junior author to work in the Divi- sion of Mollusks, United States National Museum of Natural History, Smithsonian Institution. Additional sup- port was provided in part by a Research Award from the NSF-USAP United States Antarctic Program Grant [ANTO636408] and a grant-in-aid from the Conchologists of America and the Walter E. Sage Memorial Award. We are grateful to Dr. Yu. Kantor for his many helpful and insightful comments during the course of this research, and to Roland Houart and Dr. Geerat Vermeij for their helpful reviews. The Veliger, Vol. 51, No. 1 LITERATURE CITED BEu, A. G. 2004. Marine Mollusca of oxygen isotope stages of the last 2 million years in New Zealand. Part 1: Revised generic positions and recognition of warm-water and cool-water migrants. Journal of the Royal Society of New Zealand 34(2):111-—265. CARCELLES, A. 1953. Catalogo de la malacofauna antartica argentina. Anales del Museo Nahuel Huapi 3:155—250. CASTELLANOS, Z. J. A. D. & N. LANDONI. 1993. Catalogo descriptivo de la malacofauna marina magallanica 9. Neogastropoda: Muricidae y Thaisidae. Comision de Investigaciones Cientificas de la Provincia de Buenos Aires. 26 pp. COSSMANN, M. 1903. Muricide. Essais de Paléonconchologie Comparée 5:7—66, pls. 1-3. DELL, R. K. 1990. Antarctic Mollusca, with special reference to the Fauna of the Ross Sea. The Royal Soceity of New Zealand, Bulletin 27, iv + 311 pp. HAIN, S. G. 1990. Beitrage zur Biologie der beschalten Mollusken (KI. Gastropoda und Bivalvia) des Weddell- meeres, Antarktis. Berichte zur Polarforschung 70:1—-181. HAIN, S. & P. M. ARNAUD. 1992. Notes on the reproduction of high-Antarctic molluscs from the Weddell Sea. Polar Biology 12(2):303-312. HARASEWYCH, M. G. 1984. 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D’ATTILIO. 1976. Murex Shells of the M. G. Harasewych & G. Pastorino, 2008 World, An Illustrated Guide to the Muricidae. Stanford University Press, Stanford, California; x + 284, 32 pls pp. SOLEM, A. 1972. Malacological application of Scanning Electron Microscopy, II. Radular structure and function- ing. The Veliger 14:327—336. SWOFFORD, D. L. 2002. PAUP*. Phylogenetic Analysis Using Page 10: Parsimony (*and Other Methods). Version 4.0 Sinauer Associates, Sunderland, Massachusetts. TAN, K. S. 2003. Phylogenetic analysis and taxonomy of some southern Australian and New Zealand Muricidae (Mol- lusca: Neogastropoda). Journal of Natural History 37: 911-1028. oie | wait CaN aia : , ‘1 is r i 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 References in the text should be given by the name of the author(s) followed by the date of publication: for one author (Phillips, 1981), for two authors (Phillips & Smith, 1982), and for more than two (Phillips et al., 1983). The Literature Cited section should include all (and only) references cited in the text, listed in alphabetical order by author. 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