THE NAUTILUS Volume 121, Number 1 March 28, 2007 ISSN 0028-1344 A quarterly devoted to malacology. EDITOR-IN-CHIEF Dr. José H. Leal The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 BUSINESS MANAGER Mary Jo Bunnell The Bailey-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Arthur E. Bogan North Carolina State Museum of Natural Sciences Raleigh, NC 27626 Dr. Philippe Bouchet Laboratoire de Biologie des Invertébrés Marins et Malacologie Muséum National d'Histoire Naturelle 55, rue Buffon Paris, 75005 France Dr. Robert H. Cowie Center for Conservation Research and Training University of Hawaii 3050 Maile Way, Gilmore 409 Honolulu, HI 96822 Dr. Robert T. Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. Eileen H. Jokinen 8234 E. North Shore Road Sault Ste. Marie, MI 49783 Dr. Douglas S. Jones Florida Museum of Natural History University of Florida Gainesville, FL 32611-2035 Dr. Harry G. Lee 1801 Barrs Street, Suite 500 Jacksonville, FL 32204 Dr. Charles Lydeard Biodiversity and Systematics Department of Biological Sciences University of Alabama Tuscaloosa, AL 35487 Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa P.O. Box 467 Wellington, NEW ZEALAND Dr. James H. McLean Department of Malacology Natural History Museum of Los Angeles County 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Paula M. 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Wise Department of Biology College of Charleston Charleston, SC 29424 SUBSCRIPTION INFORMATION The subscription rate per volume is US $43.00 for individuals, US $72.00 for institutions. Postage outside the United States is an additional US $5.00 for surface and US $15.00 for air mail. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 1580, Sanibel, FL 33957, USA, (239) 395- 9933! Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is published quarterly by The Bailey- Matthews Shell Museum, 3075 Sanibel-Captiva Road, Sanibel, FL 33975. Periodicals postage paid at Sanibel, FL, and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS P.O. Box 1580 Sanibel, FL 33957 NAUTILUS Volume 121, Number 1 March 28, 2007 ISSN 0028-1344 CONTENTS Richard L. Squires Paleocene pareorine turritellid gastropods from the Pacific slope of LouElla Saul NOVEM AMIETICA: joi tec aicns 1 atbrdiad te. AGBcand: deal aid. Pole Mo al ahh a eh dew, Blues Se a) dane l Brian F. Coles Vertigo malleata, a new extreme calcifuge land snail (Gastropoda: Jeff Nekola Vertiginidae) form the Atlantic and Gulf coastal plains of the USA... 2... 17 Meghna Roy Population dynamics of the fingernail clam Sphaerium occidentale D. Dudley Williams (Lewis, 1856) (Bivalvia: Sphaeriidae) in an intermittent pond ........... 29 Research Note Ilya V. Buynevich Paleoenvironmental significance of the eastern mudsnail, [lyanassa obsoleta (Say, 1822), from a microtidal coastal sequence of southern New England 37 BGTACAY oo, faethe inn coe hes hee a aA ga SNe Res, aha near Gog apa Qabed Me, iad Poa ae a aoe Mp ae 40 THE NAUTILUS 121(1):1-16, 2007 Page ] Paleocene pareorine turritellid gastropods from the Pacific slope of North Americz Richard L. Squires Department of Geological Sciences California State University Northridge, CA 91330-8266 USA richard.squires@csun.edu Angeles County LouElla R. Saul Invertebrate Paleontology Section Natural History Museum of Los 900 Exposition Boulevard Los Angeles, CA 90007 USA lousaul@earthlink.net ABSTRACT This paper presents the first detailed study of Mesalia martin- ezensis (Gabb, 1869) and Mesalia clarki (Dickerson, 1914a), the only two known pareorine (spout-bearing) turritellid gastro- pods from the Pacific slope of North America. Both species are redescribed, in light of new morphologic information that also confirms their assignment to genus Mesalia Gray, 1847, which we believe to be congeneric with Sigmesalia Finlay and Mar- wick, 1937. New stratigraphic information allows for refine- ment of the chronologic range of each species. Mesalia mar- tinezensis is of early late Paleocene (late Danian) to early late Paleocene (early Thanetian) age and ranges from northern California to northern Baja California. Mesalia clarki is of late middle to early late Paleocene age (late Selandian to early Thanetian) age and is known only fom California; in southern California it is restricted to a coralline-algal facies. Both species have considerable variability in their spiral sculpture. Mesalia originated during either the Late Cretaceous (Maas- trichtian) in northern Africa or the early Paleocene (Danian) in northern Africa and western Iran. It became widespread during the warm (greenhouse) conditions of the Paleocene and Eocene but became geographically restricted during subse- quent cooler global conditions. Mesalia is an extant genus with possibly six species, and whose total geographic range is in coastal waters in southern Portugal, southern Spain, Mediter- ranean Sea (primarily the western part), Canary Islands, and the west coast of northern Africa. INTRODUCTION The object of this study was to rectify the identification uncertainities concerning the only two known pare orie spout-bearing) turritellid gastropod species from shal- low-marine rocks on the Pacific slope of North America. They are Mesalia martinezensis (Gabb, 1869) and Mesa- lia clarki (Dickerson, 1914a). va modern workers Corresponding author: Richard Squires email: richard.squires@csun.edu (e.g., Zinsmeister, 1953) have generally placed both spe- cies in Mesalia Gray, 1847, buat. some workers (e. g., Saul, 1983a: fig. 2; Squires, 2003: table 2.4) have been ie ssitant to unequivocally use the genus name because of inad- equate morphologic information about these species. The shape of the aperture of the latter one was unknown until now, and apertural information is critical in distin- cuishing parerorine gastropods from similar looking tur- ritellids (see “Systematic Paleontology” for morphological comparisons). There has also been the possibility that Mesalia macreadyi Waring, 1914, which has commonly been put into synonymy with M. martinezensis, could be a distinct species. We conclude that there are only two species, both belonging to Mesalia, which we believe to be congeneric with Sigme salia Finlay and Marwick, 1937. In addition to providing new morphologic information about the study area Mesalia, we retine their geographical (Figure 1) and chronologic ranges (Figure 2), Both M. martinezensis and M. clarki have considerable variability in their spiral sculpture. Mesalia has long been reported (e.g., Cossmann, 1912) as originating during the Late Cretaceous in the region between northern Africa and western Iran. Our review of the literature shows that the genus most likely originated during either the Late Cretaceous (Maastrichtian) or the early Paleoce me (Danian) in this Old World region. Modem systematic studies of Mesalia are sorely lack- ing as are detailed ecologic studies of the few extant species. The classification scheme used here follows that of Bouchet et al. (2005: 249), and tase ar terms are taken from Cox (1960). Pacific slope of North America Turritella zones are taken from eel (1983b). Institutional abbreviations used in the text are: ANSP: Academy of Natural Sciences of Philadelphia; LACM: Natural History Museum of Los Angeles d ounty, Mala- cology Section; LACMIP: Natural History Museum of Los Angeles County, Invertebrate I UCMP: University of California, Berkeley, Museum of Paleontology. Paleontology Section; Page 2 THE NAUTILUS, Vol. 121, No. 1 Latitudinal Distribution of Species ]-Lower Lake 2-Martinez 3-Mt. Diablo area 4-Big Rock Creek 5-Simi Hills 6-Santa Monica Mountains 7-Mesa San Carlos California a Mesalia clarki Mesalia martinezensis Figure 1. Locales and latitudinal distribution of the study area gastropods. STRATIGRAPHY AND DEPOSITIONAL ENVIRONMENTS The geologic ages of the formations and most of the depositional environments of the formations containing the two studied species are mentioned in Squires (1997). Mesalia martinezensis is widespread in the study area (Figure 1) and always found in siltstone or sandstone be ds that formed either as storm accumulations of mol- lusk-rich assemblages in shelfal-marine depths or as lo- calized displaced shallow-marine mollusks deeper Age System Series European Mesalia Mesalia Turritella (Ma)| S¥Stemy/penies Stages | martinezensis| clarki Zones Lower = sean] | T. peninsularis Danian T. peninsularis qualeyi T. infragranulata T.1. pachecoensis Middle} Upper Lower Paleogene Paleocene Lower 65 Upper Cret.]| Maastr. Figure 2. Chronostratigraphic position of the study area taxa Ages of stage boundaries from Gradstein et al. (2004). Turri- tella zones from Saul (1983b depths. These mollusks commonly include shallow- marine mollusks such as naticid and buccinid gastropods, as well as glycymerid and crassatellid bivalves. All are indicative of normal marine salinities. The type locality of Mesalia martinezensis has been generally assigned (e.g., Keen and Bentson, 1944) to the “Martinez Formation” in the vicinity of the city of Mar- tinez, Contra Costa County, northern California (Figure 1). The history of how early workers referred to “this originally poorly defined “formation” has been summa- aed by Mallory (1959). In this present study, we follow the work of Weaver (1953), who refined the str atigraphy of the Paleocene and Eocene formations in the v icinity of the area where the “Martinez group” was first estab- lished. He established new stratigraphic names, and the rocks that pertain to the type locality of M. martinezensis belong in his Paleocene Vine Hill Sandstone. Mesalia clarki is only known from two locales: 1) its type locality (UCMP loc. 1540, see “Appendix”) on the north flank of Mount Diablo, Contra Costa County, northern California, and 2) from the Santa Monica Mountains, Los Angeles County, southern California (Figure 1). Its type locality is near the site of Stew artville, and approximately 24 km east-southeast of ihe ry oF Martinez, and, according to Dickerson (1914a: 74), this locality is “300 to 400 ft. above the base of the ates in hard, gray-green glauconitic sandstone.” Numerous mollusks have been found at this locality (Dickerson, 1914a: 75). They consist of turritellid and buccinid gas- tropods, crassatellid bivalves, and other shallow-marine species, all indicative of normal marine salinities. Turri- tella infragranulata pacheocensis Stanton, 1896, which is found at this locality, is indicative of the upper middle Paleocene (upper Selandian) (Figure 2). On the geologic map of Brabb et al. (1971), the locality plots within the glauconitic sandstone lower member of the “Martinez” Formation. Megafossils are generally scarce in the “Mar- tinez” Formation in the vicinity of this type locality (E. ), thus, it seems plausible that the fossils probably occur in storm-derived, isolated Brabb, personal commun.) lenses. Mesalia clarki in the Santa Susana Formation in the Palisades Highlands area of the Santa Monica Moun- tains, southern California, is always found near outcrops of coralline-algal limestone. Hoots (1931: 91-92, 133- 134, pL L9B) reported that these limestones are resistant, can be cliff forming, weather white, are nodular and ir- regularly bedded, up to 35 m thick, up to 1200 m in lateral extent, and commonly terminate in an abrupt wall. Additional ge ologic and/or pi aleontologic details concern- ing these limestones are mentioned in Strathearn et al. (1988), Colburn (1996), Squires and Saul (1998), Squires and Kennedy (1998), and Squires and Saul (2001). At LACMIP loc, 10508, in the Palisades Highlands area, abundant specimens of M. clarki ave found in a thin coralline-algal-rich micaceous muddy siltstone bed about | m_ stratigraphically below a blocky, coralline-algal- limestone interval approximately 24 m thick. Also fouicd R. L. Squires and L. R. Saul, 2007 Page 3 in this bed is the large neritid gastropod Corsania (Jan- uncia) rhoga Saul and Squires, 1997, as well as the bi- valves Plicatula lapidicina Squires and Saul, 1998, and Plicatula trailerensis Squires and Saul, 1998. Occurring in nearby beds in close association with the coralline- algal deposits are the gastropods Terebralia susana Squires and Kennedy, 1998, and hie abs ereenellum Hanna and Hertlein. 1939. All of these aforementioned mollusks indicate very nearshore, tropical to subtropical conditions (Squires and Saul, 1997; Squires and Kennedy, 1998). The latter workers concluded that the coralline-algal limestones, like those at locality 10508, were deposited in a protected bay (no deeper than 40 to 70 m) with warm-algal-limestone buildups associated with shoals on the bay floor. These limestone buildups are very similar in lithology and sedimentologic/tectonic setting to limestones in the Paleocene Sepultura and Ba- hia Ballenas formations in northern Baja California (Ab- bott et al., 1995), as well as similar to limestones in the upper Paleocene to lower middle Eocene Sierra Blanca Limestone in Santa Barbara County, southern California. These limestones were deposited when tectonic plate- edge strain in the fore-arc basin caused local basement highs to form within the otherwise deeper marine envi- ronment (Whidden et al., 1995; Abbott et al., 1995). It is likely that the Santa Susana Formation coralline-algal limestones formed under similar conditions. Although Mesalia clarki and Mesalia martinezensis hoe occur in the Santa Susana Formation in the Santa Monica Mountains, southern Califormia, they never oc- cur together in the same beds. Mesalia martinezensis is not associated with the coralline-algal facies there or any- where else in the study area. PALEOBIOGEOGRAPHIC IMPLICATIONS Kiel and Bandel (2004: 120, fig. 71) reported two speci- mens of Mesalia cf. multilineata (J. Miiller, 1851) from Cenomanian strata at the Kasssenberg quarry in Ger- many. If these specimens actually belong to Mesalia, they would be the geologically oldest. The conical-turriculate shell with convex whorls bearing strong spiral ribs does resemble Mesalia, but there are no specimens with an intact aperture nor with a protoconch. Turritella multi- lineata J. Miiller (1851: 29, pl. 4, figs. 4, 6) was originally reported from the lower Campanian Aachen strata of Germany, thus, it is considerably younger than the Kass- senberg quarry material. Turrite lla mattiline cata Was also figured by G. Miiller (1898: 97, pl. 13, figs. 4, 5), who reported it from middle Santonian to lower Campanian strata at Braunschweig/Ilsede, Germany. There is also a mention of T. multilineata in Kollmann and Odin (2001: 441), and they also consider this Campanian species to belong to genus Turritella. The pleural angle of J. Miiller’s figure is much narrower than that of Kiel and Bandel (2004), and in our opinion, Kiel and Bandel’s M. cf. M. multilineata (J. Miiller) does not look like J. Miiller’s species. Kiel and Bandel (2004) tentatively syn- onymized ]. Miiller’s specimens and G. Miiller’s speci- mens with their Kassenberg quarry specimens. We be- lieve, however, that Kiel Gnd Bandel’s Cenomanian specimens have nothing to do with T. multilineata and represent, at best, a very questionable Mesalia. More specimens of this possible Mesalia from the Cenomanian of Germany are needed in order to determine its generic assignment. Cossmann (1912: 126) reported the chronologic range of Mesalia to be Late Cretaceous (Turonian) to Recent, as did Wenz (1939), who apparently simply reiterated Cossmann’s findings. We were unable, however, to cor- roborate any of Cossmann’s Cretaceous occurrences. He reported Mesalia gazellensis Whitfield (191; 424, pl. 9, fig. 10) as being from the Turonian of Syria, but the aperture of his, species is unknown. In addition, the sculpture is obsolete, which is unlike Mesalia. Cossmann (1912: 126) listed five Mesalia species of Late Cretaceous (Senonian) age, and these are discussed in the following sentences. Arcotia indica Stoliczka (1868: 215, 469, pl. 16, figs. 12, 12a; pl. 19, fig 6) from southern India is not a Mesalia. This species is also dis- cussed later under “Systematic —. Specimens of Turritella ventricosa Forbes (1846: 123, pl. 13, fig. 3; Stoliczka, 1868: 227, pl. 17, fig. 15) ‘ne southern Indi ia are missing the aperture. Turritella martinezensis Gabb (1869: 159. pl. 28, fig. 51) from California is not of Cre- taceous age. Mesalia nettoana White (1887: 164-165, p 18, figs. 3, 4) from the Maria Farinha beds in Brazil is oe age. Mesalia hebe White (1887: 165, pl. 1S, fig. ), also from Brazil, looks like a juvenile specimen of M. ne eer Cossmann (1912: 126) also listed two Late Cretaceous (Maastrichtian) species. The first one is Mesalia jovisam- monis (Quaas, 1902: 256, pl. 26, figs. 1S—20), which Quaas reported, in a very generalized way, to be associ- ated with the Exogyra overwegi biozone at Ammonite Hill in the Great Sand Sea in western Egypt. This bio- zone can also be recognized in the Maastrichtian (but not latest Maastrichtian) part of the Ammonite Hill Member of the Campanian to Paleocene Dakhla Formation in western Egypt (Barthel and Herrmann-Degen, 1981). Tantawy et al. (2001) assigned this member an early to early late Maastrichtian age, based on planktic forami- nifera, calcareous siannofossils , and macrofossils. They also determined, however, that the entire formation ranges in age from early Maastrichtian to early Danian. Immediately above the widespread K/T disconformity in the region, a sedimentologically complex sequence marks the lower Danian Bir Abu Minquar horizon, which contains a mixture of Maastrichtian (reworked) and Da- nian fossils, including both microfossil and macrofossil species (e.g., including some ammonites.). Unfortu- nately, Quaas did not “provide any information as to where actly in the stratigraphic section he collected the specimens of M. jovisammonis. His specimens were lost, so it is not possible to match their rock matrix to actué ul outcrops. Recollecting of this gastropod is necessary to decipher its exact geologic age. Page 4 THE NAUTILUS, Vol. 121, No. 1 14 Figures 3-15. Type species of Mesalia and Sigmesalia, plus comparative pictures of Mesalia solida (Deshayes, 1861). Specimens coated with ammonium chloride, 3-9. Mesalia mesal (Adanson, 1757), Baie de Hann, Senegal, West Africa (Recent). 3-7. Hypotype LACM 173163. 3. Apertural view, height 45 mm, diameter 15 mm. 4. Tip of specimen shown in previous figure, height 14 mm, diameter 5.5 mm. 5. Protoconch and earliest spire whorls, apertural/ right-lateral view, height 1 mm, diameter 0.7 mm. 6. Base, diameter 15.6 mm. 7. Abapertural view, height 45 mm, diameter 15 mm. 8-9. Hypotype LACM 173164. 8. Oblique apertural view, height 51 mm, diameter 16.3 mm. 9. Close-up of abapertural view of last whorl, diameter 17.2 mm. 10-13. Mesalia koeneni (Le Renard, 1994), LACMIP hypotype 13397, Grignon, Paris Basin, France (middle Eocene, Lutetian Stage), height 45.8 mm, diameter 20.3 mm. 10. Apertural view. 11. Close-up of aperture. 12. Right-lateral view (outer lip broken), 13. Base. 14-15. Mesalia solida (Deshayes, 1561), hypotype LACMIP 13398, Le Guépelle, Paris Basin, France (late Eocene). 14. Apertural view, height 21.6 min, diameter S.1 mm. 15. Protoconch and earliest spire whorls, apertural to slightly right-lateral view, height 1 mm, diameter 0.7 Tim. R. L. Squires and L. R. Saul, 2007 Page 5 Abbass (1963: 39-40, pl. 2, figs. 20-22) illustrated M. jovisammonis from eastem Egypt and referred to it as Mesalia (Woodsalia) jovisammonis of Maastrichtian age. He did not provide, however, any discussion as to how this age was determined. Mesalia cf. M. multisulcata (Lamarck 1804), ea by Serra (1937: 313-315, pl. 16, figs. 12, 12a, 13) from near Tripoli, Libya, looks like it might be conspecific with Mesalia jovisammonis. Serra provided no detailed stratigraphic or geologic age information. The second species that Cossmann (1912) listed from the Maastrichtian is Mesalia fasciata (Lamarck, 1804) from Iran. Cossmann based this occurrence on work by Douvillé (1904: 329-330, pl. 47, figs. 23-27), who re- ported M. fasciata from the * ‘Couches: a Cérithes” beds in the Luristan region in west-central Iran. Douvillé (1904) believed that these Iranian specimens of M. fasciata, whose type locality is in middle Eocene (Lutetian Stage) strata at Grignon in Paris Basin, France (Eames, 1952: 34), are of Maastrichtian age, but the “Couches a Cérithes” beds contain the bivalve “Cardita” beaumonti dArchiac and Haime, 1854, which is diagnostic of earli- est Danian age in Iran and Pakistan (Douvillé, 1928: Eames, 1952; Davies, 1975). Mesalia fasciata is long- ranged geologically (early Paleocene to late Eocene) and widespread geographically (western Europe to Pakistan) (Eames, 1952). Another Mesalia that needs investigation as to its pre- cise stratigraphic position and geologic age is Mesalia foucheri Pervinquiére (1912: 44, pl. 3, figs. 6-15), from the Ghadames (Garat Temblili) region in Tunisia, north- ern Africa. Pervinquiére (1912: 336) reported the species as being of Maastricthian age, but no critical geologic details are provided. He did differentiate between Maas- trichtian and Danian fossils; thus, like in nearby Egypt and Libya, the stratigraphic section containing M. foucheri and other macrofossils in Tunisia, also spans the K/T boundary. Two species of so-called Woodsalia Olsson, 1929, from Upper Cretaceous (Campanian?) strata in northwestern Peru (Olsson, 1944) might eventually be placed in Me- salia, once their apertures become known. They oo Woodsalia paitana Olsson (1944: 69-70, pl. 11, fig. 5) and W ‘oodsalia paitana robusta Olsson (1944: 70, pl. 11, figs. 3, 9). The so-called Mesalia (Mesalia) mauryae Allison (1955: 414415, pl. 41, fig. 3; Perrilliat, 1989: 149, fig. 51h) from the upper Aptian upper member of the Alisi- tos Formation, Punta China region, Baja California, Mexico, is, according to Squires and Saul (2006), Turri- tella seriatimgranulata Roemer, 1849. In addition to the above-mentioned Old World Danian species of Mesalia, three New World Danian species are known from the Gulf Coast of the southeastern United States. They are from the Clayton Formation (Palmer and Brann, 1966), which is of earliest Danian age (Dock- ery, 1986). The three species are: Mesalia silenkene nsis (Aldrich, 1894: 246-247, pl. 13, figs. 4a, 4b, 6; Stenzel and Turner, 1942: card 110); Mesalia hardemanensis (Gabb, 1860: 392, pl. 68, fig. 15; Stenzel and Turner, 1942: card 116); and Mesalia pumila (Gabb, 1860: 392 pl. 68, fig. 14; Stenzel and Turner, 1942: card 118). In suminary, our search of the bieranare revealed that Mesalia most likely originated during either the Maas- trichtian in northern Africa or the early Paleocene (Da- nian) in northern Africa and western Iran. During the Danian it spread quickly to the Gulf Coast of the Wned States by means of westward-flowing ocean currents emanating from the western Tethyan region. These cur- rents, which existed during the Late Cretaceous (Gor- don, 1973; Johnson, 1999) aad continued into the Paleo- cene and Eocene (Saul, 1986; Squires, 1987), were part of a circumglobal-tropical current that contributed to a widespread dispersal of marine biota (Haq, 1981). By the late Danian, it reached California and northern Baja California, Mexico, as well as Belgium (Cox, 1930; Glib- ert, 1973). By middle Paleocene, it reached Greenland (Kollmann and Peel, 1983), and by the late Paleocene, it reached Nigeria (Adegoke, 1977). During the Paleocene and Eocene, Mesalia reached its peak div ersity and be- came most widespread, with occurrences mainly in the Old World western Tethys Sea region. We did not de- tect, however, any reported occurrences in Australia, New Zealand, Japan, or Antarctica. The Paris Basin of France (see Cossmann and Pissarro, 1910-1913), south- western Nigeria (Adegoke, 1977), and the Gulf Coast of the United States (Stenzel and Turner, 1940, 1942: Palmer and Brann, 1966) are the principal areas in which numerous species of Mesalia have been anlage Some species became very widespread. For example, a mentioned earlier, Mesalia fasciata ranged trom ae Paris Basin, France to Pakistan (Eames, 1952). After the warm greenhouse conditions that existed during the Eocene, Mesalia was much reduced in its distribution and mainly occurred in what is now the Mediterranean Sea region (Cossmann, 1912). MODERN MESALIA Mesalia is extant and review of the scant literature, as well as use of the intemet (note: was particularly helpful), revealed as many as possibly six species. The they are the ae M. mesal (Adanson, 1757), M. brevialis Lamarck, 829. M. varia Kiener, 1843; M. opalina Adams and See 1850; M. freytagi von Maltzan, 1884; and M. flammifera Locard, 1897, which includes the subspecies M. flam- mifera flammifera Locard, 1897 and M. flammifera sim- plex Locard, 1897. There is much confusion as to exactly how many species there are, and potential synonyms need to be resolved. For example, some workers (e. o., Smith, 1915; Bowles, 1939) equated M. mesal with M. brevialis, but other workers (e.g., Advovini and Cos- signani, 2004) separated them. Bowles (1939) gave a thorough review of the nomenclatural history of Mesalia brevialis. Page 6 THE NAUTILUS, Vol. 121, No. 1 A comprehensive malacological study of the modern species of Mesalia is greatly needed. Because of the un- certainties stemming from the poorly y known sy! stematics, it is confusing to try to determine which species is found where. We were able to establish with certainty (see references below), however, that modern Mesalia is only found in the Atlantic coastal areas of southern Portugal, southwestern Spain, Morocco, Canary Islands, Western Sahara, Mauritania, Senegal, and Guinea, as well as in the westernmost RicAnereanean Sea, particularly in the Alboran Sea (i.e., Strait of Gibraltar to southerm Spain on the north and Morocco on the south) and the Aegean Sea region of western Turkey. Mesalia mesal and M. brevialis have the widest distri- bution. Mesalia mesal has been reported from the Al- garve region of southern Portugal, the Algeciras region of southwestern Spain, and the “‘Alboran Sea ( (Poppe and Goto, 1991), the Canary Islands (Macedo and Borges, 1999), Senegal (Bouchet, 1977: fe nae and Cossignani, 2004), and western Turkey (Demir, 2003). Me salia bre- vialis has been reported from the Algarve region of southern Portugal (Afonso et al., 2000; Alves et al., 2003), southwestern Spain and the Alboran Sea (Hidalgo, 1917), Senegal (Ardovini and Cossignani, 2004), and Guinea (Pasteur-Humbert, 1962). Mesalia opalina has been reported from the Canary Islands and Morocco (Poppe and Goto, 1991), as well as from Mauritania (Ar- dovini and Cossignani, 2004). The other modern species/ subspecies of Mesalia are apparently restricted to the northwestern coast of Africa (Ardovini and Cossignani, 2004). Mesalia melanioides Reeve, 1849, was reported (Smith, 1915) to be from West Australia, but this species is now the type species of Neodiastoma Cotton, 1932 which differs from Mesalia by having axial sculpture on the early spire. Marwick ( 1957) summarized the system- atics of Neodiastoma and classified it as a pareorine. Mesalia is found today on both muddy and sandy bot- toms in coastal waters ranging in depth from lower in- tertidal to 20 m (Hidalgo, 1917; Pasteur-Humbert, 1962: Poppe and Goto, 1991; Afonso et al., 2000; Demir, 2003; Alves et al., 2003). Bouchet (1977) reported that M. me- sal, although not common there, can be found in the seaward part of mz WgrOve-sw, saad systems along the coast of Senegal. Specimen LACM 17 316 ( (Figures 3 3-7) of M. mesal was collected in pees 5m depth, on sand and rubble in Senegal. Large numbers of M. mesal have been reported \fonso et al., 2000) as almost always being partially in- faunal (with their apices pointed upward) when found on exposed low-tide mud flats on the inner lagoon sides of islands within the Rio Formosa coastal-lagoon system of southern Portugal. The modern ecological parameters mentioned above are not totally reliable for fossil Mesalia because prefer- ences for substrate and depth of water might have pos- sibly changed over time. In addition, the fossil occur- rences of Mesalia had a pan-Tethyan distribution, whereas the modern occurrences have contracted pri- marily to the Iberian Peninsula, Alboran Sea, and north- western Africa. SYSTEMATIC PALEONTOLOGY Superfamily Cerithioidea Fleming, 1522 Family Turritellidae Lovén, 1S47 Discussion: Allmon (1996: 9-12, table 1) thoroughly reviewed the history of the dlasaheaton of ‘umitellid gastropods and listed the five subfamilies and all the genera/subgenera within each subfamily. These subfami- lies are: Turritellinae Loven, 1847; Protominae Marwick, 1957; Pareorinae Finlay and Manwick, 1937: Vermicu- lariinae Lamarck, 1799; and Turritellopsinae Marwick, 1957. Bouchet et al. (2005) included the first four of these subfamilies, but removed Turritellopsinae. Instead, they included subfamily Orectospirinae Habe, 1955. Subfamily Pareorinae Finlay and Marwick, 1937 Discussion: Pareorine turritellids are characterized from the other subfamilies of family Turritellidae by hav- ing an aperture obliquely effuse over the anterior end of the columella and forming a sinus (short spout), with the adapical margin of the sinus usually making a spiral ridge on the eolamells | Marwick, 1957). Mesalia can be readily identified if its aperture is in- tact, but when it is missing, workers have commonly misassigned it to the similar looking genus “Twrritella” Lamarck 1799, sensu lato, a group comprising at least 35 genera and subgenera names (Allmon, 1996), all of which are turritellines whose apertures do not have a sinus (short spout) at the anterior end of the aperture nor have a spiral ridge on the columella. In addition, according to Smith (1915), the cormeous operculum of Mesalia is pau- cispiral and not multispiral, as in “Turritella,” but this later distinction is not useful when studying fossil spe- cles. Ten pareorine genera were listed by Marwick (1957), who also provided i illustrations of the growth- line traces of some of these genera. Comparative information about the str: itigraphic range, grow th-line details, whorl protile, and protoconch shape of a of these genera was given by Allmon (1996; table 1 Genus Mesalia Gray (nomen nudum, 1840), 1847 Type Species: Cerithiwm mesal Adanson, 1757 [=Tur- ritella mesal Deshayes, 1843], by original designation; Recent, southern Portugal, southwestern Spain, Alboran Sea, Canary Islands, Seneval, and western Turkey. Description: Small to large (wp to approximately 95 mm shell height), turritelliform, slender to conical ro- tund. Pleural angle ranges from 15° to 41°. Protoconch conical, small, smooth, and approximately two whorls. Protoconch to teleoconch transition gradual. Teleoconch whorls up to 16, whorl sides convex to flattish/concave. Sculpture on early juvenile teleoconch whorls variable, ranging from ne varly smooth or with very fine, unicostate, bicostate, or tricostate spiral lirae; sculpture on adult R. L. Squires and L. R. Saul, 2007 Page 7 whorls highly variable, ranging from smooth to numer- ous, weak to moderately strong closely spaced spiral ribs, but less commonly with fewer and more prominent spiral ribs. Growth lines parasigmoidal on last whorl (including iene, lateral sinus variable in amount of concavity (fex- ure). Aperture with shallow effuse spout, ranging from cee constrained to broad. Adapical edge of spout usually forms weak spiral ridge that continues onto col- umella. Discussion: Mesaliopsis Thiele, 1929 [type species: Mesalia opalina (Adams and Reeve, 1850)], Recent, was reported by Wenz (1939) to be a subgenus of Mesalia, but future work ah show it to be synonymous with Mesalia. Mesalia somewhat resembles Lithotrochus Conrad, 1855, of Jurassic age from Chile, South America. Coss- mann (1912: 125) reported Lithotrochus to be a junior synonym of Mesalia, but Wenz (1938: 280, fig. 596) and Cox (1960: 1245-1249, fig. 159,11) believed Lehatroe hus to be a trochid. It is an extraordinarly large gastropod (height 125 mm) with a wide pleural angle, deme d upper spire, turritelliform shape, anteriorly carinate whorls, and relatively few spiral ribs. Details of its aperture are un- known. Cossmann (1912: 125) also reported Lithotrochus to be a junior synonym of Arcotia Stoliczka, 1868, whose OPE species, Arcotia indica Stoliczka (1868: 215, 469, pl. 16, figs. 12, 12a; pl. 19, fig. 6) is from Upper Cretaceous (Trichinopoly Group) strata near the town of Alundan- apooram, southern India. According to Sundaram et al. (2001; fig. 3), this town’s name is also referred to as Alundalippur and, from information in their map, this town is underlain by the Kulakkalnattam Formation of Turonian age. Wenz (1939) synonymized Arcotia with Mesalia. Finlay and Manwick (1937) reviewed the mor- phology of Arcotia and reported that, based on_ its straight growth lines and open umbilicus, this genus is not a synonym of Mesalia. They reported, furthermore, that Aroctia appears to be a mathildid. Bandel (2000) came to the same conclusion. Mesalia is similar to the pareorine Woodsalia Olsson 1929. whose type species, Woodsalia negritosensis Ols- son (1929: 13-15, pl. 4, figs. 5, 6) is from lower Eocene rocks in northwestern Peru. Woods (1922: 78-79, pl. 7 figs. 5-7; pl. 8, figs. 1-3) and Wenz (1939: 651, fig. 1852, two views) also illustrated this species. The full shape of the aperture of this Peruvian gastropod, however, is not known. Genus Sigmesalia Finlay and Marwick, 1937, was originally erected to accommodate a group of Eocene gastropods from the Paris Basin, France that were pre- viously identified as Mesalia. There has been no consen- sus as to whether or not Sigmesalia is a distinct genus. Marwick (1957) reported it to be a separate genus, as did Le Renard (1994). Eames (1952) reported it to be a subgenus of Mesalia, and Palmer and Brann (1966) re- ported Sigmesalia to be synonymous with Mesalia. Various views of representative specimens of the type species of Mesalia are illustrated in Figures 3-9, and various views of a representative specimen of the type species of Sigmesalia are illustrated in Figures 10-13. Its type species, Sigmesalia koeneni Le Renard, 1994 [new name for Turritella sulcata Lamarck, 1804 (original des- ignation), non Bosc, 1801], is of middle Eocene (Lu- tetian) age and from Paris Basin, France. Finlay and Marwick (1937) stated that the aperture and growth lines of the type species of Mesalia seem to be generically different than those of the Paris Basin shells, but the ry did not provide any details. Davies (1971: 312, figs. 677a, 677b) mentioned that the growth lines of Sigmesalia have a more flexed outer lip sinus than does Mesalia. According to Marwick (1957: 163), Sigmesalia differs from Mesalia by usually having a wider pleural angle. The following paragraphs deal with our observations con- cerning these proposed diagnostic features of Sigmesalia. Inspection of representative specimens of several of the Eocene Paris Basin species, including the type spe- cies of Sigmesalia, stored in the LACMIP collection, as well as inspection of photographs of 17 Paris Basin spe- cies (see Cossmann and Pissarro, 1910-1913: pl. 21, figs. 126-1 to 126-15), revealed variability in the shape of the aperture of Sigmesalia. For example, the aperture of Sig- mesalia incerta (Deshayes, 1832; Cossmann and Pissarro, 1910-1913: pl. 21: fig. 126-4, two views) is similar to that of M. mesal, in that the spout is broad and not well constrained. The aperture of Sigmesalia koeneni how- ever, is better developed (Cossmann and Pissarro, 1910— 1913: pl. 21, fig. 126-15). The amount of flexure of the outer lip sinus is basically similar in all the Eocene Paris Basin species, although Mesalia solida (Deshayes, 1861) does show some vari- ability. The amount of flexure of this feature is variable in M. mesal and can be similar to the amount seen on Eocene Paris Basin species. Variability in the amount of flexure for both the Eocene and modem Mesalia shells can also occur in proximity of growth checks and break- ages of the outer lip incurred during the life of the gas- tropod. The pleural angle of the Eocene Paris Basin shells is quite variable, ranging from 21° to 41°, but the low end of this range [e.g., Mesalia ecki (Laubriére, 1881; Coss- mann and Pissarro, 1910-1913: pl. 21, fig. 126-2)] is very close to the value (16° to 18°) on M. mesal. Sigmesalia koeneni has one of widest pleural angles (41°; see Figures 10 and 12). For meee purposes, an illustration (Figure 14) is provided for Sigme salia solida. It has a relatively narrow pleural angle of 25°, more like that found on M. mesal (Figures 3, 4, and 7) Other morphologic features that are variable on the Eocene Paris Basin shells are strength and number of spiral ribs, pattern of development of sculpture on the early juvenile teleoconch whorls, and degree of indenta- tion of the suture. Even the strength of "he spiral ridge (Figure 10) on the columella is variable. Mesalia mesal pos- sesses a spiral ridge on the columella, as do most specimens of Sigmesalia koeneni (compare Figures 8 and 11) Page 8 THE NAUTILUS, Vol. 121, No. 1 A few species of Sigmesalia are similar to Mesalia in having a relatively narrow pleural angle and bicostate sculpture on the ‘early juvenile where but not on the adult whorls. They are the following: Sigmesalia instabi- lis (Briart and Cornet, 1873: 86, pl. 12, figs. 9a-9b) of early Paleocene (Danian) age from Belgium; Sigmesalia salvani (Adegoke, 1977: 86-88, pl. 14, figs. 10-16) of Paleocene age from Nigeria; and Sigmesalia fasciata (Lamarck, 1804: 217) from Eocene strata in France, Bel- gium, and Pakistan (Cossmann and Pisarrro, 1910-1913: Cox, 1930; Eames, 1952); Sigmesalia pagoda ( (Cox, 1930: 160-161, pl. 18, figs. Ga—b, 7a—b) from Eocene strata in Pakistan; Sigmesalia biplicata (Bowles, 1939: 328, pl. 34, figs. 6, 8) from Paleocene strata in Alabama; and Sigme- salia gomin (Bowles, 1939: 326-327, pl. 33, fig. 9) from Paleocene strata in South Carolina. The protoconchs of Mesalia mesal and Sigmesalia solida are very similar (compare Figures 5 and 15); both are small, smooth, have essentially tie same shape, and the transition to the teleoconch is gradual. In summary, we found that the morphologic features of the Eocene Paris Basin shells are variable. We could find no reliable, constant morphologic characters to dis- tinguish Mesalia from Sigmesalia; hence, we regard them as congeneric. Mesalia martinezensis (Gabb, 1869) (Figures 16-23) a martinezensis Gabb, 1869: 169-170, 228, pl. 25, fig. 1; Dickerson, 1914a: pl. 13, fig. 10; Waring, 1917: 100, pl. 7 fig. 5. Turritella maccreadyi Waring, wes 783; Waring, 1915: fig. [not fig. 14]; Waring 1917: S7-SS, pl. 12, fig. 10. Mesalia maccreadyi (W ering), uedes-Mejia, 1989: 176-177, pl. 3, figs. 3-6. Mesalia martinezensis (Gabb).—Cossmann, 1912: 126; Stew- art, 1927: 353-354, pl. 25, fig. 1; Schenck and Keen, 1940: pl. 20, fig. 5; Merriam, 1941: 127-128, pl. 39, figs. 1-5, 7; Zinsmeister, 1974: 11S-119, pl. 12, figs. 5, 9; Zinsmeister, 1983: pl. 2. fig. 14: Paredes-Mejia, 1989: 173-176, pl. 3, figs. 7-LO; Saul, 1983a: text-fig. 2, pl. 1, fig. 2. Mesalia clarki (Dickerson).—Zinsmeister, 1983: pl. 2, fig. o 13. Description: Large (up to approximately 95 mm height). Tarritelliform: Pleural angle approximately 20°. Protoconeh unknown. Teleoconch up to 12 whorls, in- creasing rapidly in size from the apex. Suture slightly impressed. Sculpture consisting only of spiral ribs of dif- fering strength but dominate d by carina located anteri- orly; ribs generally becoming stronge r with growth; spiral threads on inte rspaces and on carina surface, Carina usu- ally strongly angulate but can be rounded or even sub- dued. Posterior to carina, several widely spaced spiri al ribs of variable strength occur, ranging from tertiaries to primaries: two ribs on uppermost spire, three to four on upper spire and one to three on lower spire. Anterior to carina, several spiral ribs of variable strength occur, rang- ing from tertiaries to primaries: approximately five ribs On uppel spire and one to two ribs (both occasionally quite prominent) on lower spire. Last whorl with three primaries, both posterior and anterior to carina. Base with three secondaries, anteriormost one weak: ribs ob- solete on short neck. Aperture relatively small, D- shaped; columella relatively broad, smooth; spout effuse and short with anterior end projecting slightly; growth- line trace of last whorl (including base) par asigmoidal, with lateral sinus flexure strongest in vicinity of carina. Holotype: mim. ANSP 4344, height 57 mm, diameter 23 Type Locality: not given). Martinez, northern California (details Geologic Range: Late early Paleocene to early late Paleocene (near the Danian-Selandian bounde ury to early Thanetian), Distribution: DANIAN = Turritella peninsularis qua- leyi Zone: lower San Francisquito Formation, Warm Springs Mountain, Los Angeles County, southern Cali- fornia (new stratigraphic occurrence, LACMIP. loc. 21581). NEAR THE DANIAN-SELANDIAN BOUND- ARY = Turritella peninsularis qualeyi Zone transitional with Turritella peninsularis Zone: Martinez Formation, Herndon Creek east of Lower Lake, Lake County, north- ern California (Stanton, 1896 [faunal list]; Dickerson, 1914a; Merriam, 1941); upper Las Virgenes Sandstone, Simi Hills, Ventura County, southern California (Waring, 1917; Nelson, 1925 [faunal list]; Merriam, 1941; Zins- meister, 1983; Saul, 1983a). PROBABLY NEAR THE DANIAN-SELANDIAN BOUNDARY: Reworked specimens in Santa Susana Formation, Poison Oak Can- yon, north side Simi Valley, Los Angeles County, south- erm California (new stratigraphic occurrence, LACMIP loc. 21554); Reworked specimens in Stokes Canyon Breccia Member of the middle Miocene Calabasas For- mation, Stokes Canyon, Santa Monica Mountains, Ven- tura County (new stratigraphic occurrence, LACMIP loc. 25281). SELANDIAN = Turritella peninsularis Zone: Lower Vine Hill Sandstone, Martinez area, Contra Costa County, northern California (Weaver, 1953 [faunal list]); lower San Fri mcisquito Formation, Finvou Ridge east of Big Rock Creek, Valymero area, Ante lope Vv alley, Los Ange sles County, southern California (Dichensen: 1914b anal list]; Merriam, 1941; Kooser, 1980 [faunal list]); lower Santa Susana Formation (= “Martinez ma- rine member” of Nelson, 1925 [faunal list]), Simi Hills, Ventura County, southern California (Kew, 1923 [faunal list]; Nelson, 1925 [faunal list]; Zinsmeister, 1983; Saul, 1983a). LOWER THANETIAN = Turritella infragranu- lata Zone: Upper Vine Hill Sandstone, Martinez area, Contr re . County, northern California (Weaver, 1953 ; upper Santa Susana Formation, Palisade 2S Highl: ads. inta Monica Mountains, Los Angeles County, southern California (new stratigraphic occur- rence, LACMIP locs. 7060 and 11717); Sepultura For- mation, Mesa San Carlos, northern Baja California, Mexico (Paredes-Mejia, 1989), R. L. Squires and L. R. Saul, 2007 Page 9 Figures 16-23. Discussion: The largest specimens of this species oc- cur in the lower San Francisquito Formation, Pinyon Ridge east of Big Rock Creek, Valymero area, Antelope Valley, Los Angeles County, southern California. There is considerable variability in the strength of the spiral ribs on M. martinezensis. Most specimens are Cari- nate on all whorls, including the last whorl. On some specimens. however, the carina becomes weaker on the later whorls as the other spiral ribs become stronger, giving these whorls a convex shape (Figures 19-21), like Mesalia martinezensis (Gabb, 1869). Specimens coated with ammonium chloride. 16-18. Hypotype LACMIP 13399, height 55 mm, diameter 25.4 mm. 16. Apertural view. 17. Oblique apertural view. 18. Abapertural view. 19. Hypotype LACMIP 13400, LACMIP loc. 22557, apertural view, height 36.3 mm, diameter 14.1 mm. 20. Bhai LACMIP 13401, LACMIP loc. 21607, abapertural view, height 38.3 mm, diameter 19.3 mm. 21. Hypotype LACMIP 13402, LACMIP loc. 22698, abapertural view, height 33.9 mm, diameter 21.6 mm. 22. Hypotype LACMIP 13403, LACMIP loc. 26897, apertural view, height 10.6 mm, diameter 6.5 mm. 23. Hypotype LACMIP 13404, LACMIP loc. 22330, base, diameter 17.2 mm. shells misidentified by some workers as Turritella ma- creadyi Waring, 1914. The overall felecuantel morphology of the 10 mm-high tip of Mesalia martinezensis superficially resembles that of the 15-mm high mathildid Carinathilda diminuata (Perrilliat, Vega, aaa Corona, 2004) illustrated by Kiel et al. (2002: 329-330, fig. 2.4) from the lower Maastrichtian of the Mexcala Formation, Guerrero, southern Mexico. Carinathilda diminuata is definitely a mathildid because it has a heterostrophic protoconch. The resemblance be- Page 10 THE NAUTILUS, Vol. 121, No. 1 tween these two gastropods, nevertheless, provides evi- dence that the Late Cretaceous mathildids and lower Paleogene turritellids can have similar looking adult shells. Mesalia martinezensis resembles “Mesalia” virginiae Stilwell et al. (2004: 29-30, pl. 5, figs. 6-10) from lower Paleocene (Danian) rocks on Seymour Island, Antarctic Peninsula, but M. martinezensis has a subtle effuse spout rather than the longer and more distinct, twisted narrow anterior canal that “M.” virginiae possesses. In addition, M. martinezensis has stronger ribs and a parasigmoidal growth line, rather than an opisthocyrt one on the last whorl. In our opinion, the aperture of “M.” virginiae is unlike that of Mesalia. Gabb (1869) mentioned that the broadly expanding whorl of martinezensis approaches that seen on Turri- tella robusta Gabb (1864: 135, pl. 21, fig. 74; not = T. (Haustator) robusta Grzybowski, 1899), but Merriam (1941: 128) reported that the Late Cretaceous T. ro- busta, which is represented by a single poorly preserved type specimen, is probably not a Me salia. This type speci- men has an umbilicus, therefore it is not a turritellid. It is from the Redding area, northern California, and not from Tuscan Springs, as erroneously reported by Mer- riam (1941). Jones et al. (1975: pl. 1, fig. 19) identified this specimen, which is of Turonian age, to be Glauco- nia? robusta (Gabb, 1864). Merriam (1941: 10, 116) stated that mainly in profile the Pacific slope Miocene Turritella temblorenesis Wiedey, 1928, might readily be confused with Mesalia martinezensis. The latter also resembles the Pacific slope Miocene Turritella temblorensis tritschi Hertlein, 1928, and Turritella ocoyana Conrad, 1857. The latter, how- ever, has a different growth line. In addition, T. martin- ezensis strongly resembles Turritella fredeai Hodson, 1926, of Miocene age from northern Colombia and northern Venezuela. None of these above-mentioned Miocene species, however, has the effuse spout of Me- salia. Mesalia clarki (Dickerson, 1914a) Mores 24-39 (Figures 24—32) Mesalia clarki (Dickerson).—Merriam, 1941; 128, pl. 39, fig. 6 Zinsmeister, 1983: table 1, pl. 2, fig. 14 Turitella {sic| clarki Dickerson, 1914a: 142-143, pl. 13, fig. 8. Description: mm height). Turritelliform. Pleural angle approximately 21 to 22°. 12 whorls, consisting of two whorl shapes: flatish rounded Medium small (up to approximately 31 Protoconch unknown. Teleoconch up to and anteriorly aneulate. Sutural area indente - Sculpture consisting only of spiral ribs, variable in number, strength, and spacing. Flattish to rounded shen shape: upper spire aah one or two secondaries on posterior part and two (bicostate) primaries on anterior part; lower spire and last whorl with seven to eight nearly equal strength primaries (anteriorly located ribs can be some- what ‘angulate); spiral threads on all interspaces; poste- riormost rib part of broad band; ribs on anterior part of whorl tend to be slightly stronger than posteriorly located ribs: base of last whorl with thee ribs. Angulate whorl shape: upper spire with one secondary on posterior part and two (bicostate), well dev eloped, f lat-topped prima- ries on anterior part; lower spire and last whorl with three primaries on posterior part and two stronger pri- maries, with one secondary in between each, on anterior part: spiral pas on all interspaces. Base (including short neck) of last whorl with approximately seven, evenly ied ies interspaces and ribs covered by spiral threads. Aperture small; columella narrow with thin cal- lus, occasionally with single, weak fold, slight twist on anterior end of columella. Spout effuse, short, and nar- row. Growth-line trace on last whorl (including base) parasigmoidal, with lateral sinus flexure strongest medi- ally. Holotype: UCMP 11936, height 25 mm, diameter 16.5 mm. Type Locality: UCMP loc. 1540 Geologic Range: Late Paleocene = Turritella infra- granulata Zone. Distribution: “Martinez” Formation, northeast side of Mount Diablo, Contra Costa County, northern Califor- nia (Dickerson, 1914a; Merriam, 1941; Zinsmeister and Paredes-Mejia, 1955 [faunal list]; upper Santa Susana Formation, Trailer and Quarry canyons, Los Angeles County, Santa Monica Mountains, southern California (Strathearn et al., 1988 [faunal list]; Squires and Saul, 1998: 1025). Discussion: Mesalia clarki is abundant in the upper Santa Susana Formation at LACMIP loc. 10508, in the Santa Monica Mountains, Los Angeles County, southern California. The anterior ends of the shells are very frag- ile, and nearly all the specimens have incomplete aper- tures. None of the specime ns has retained their proto- conch, and most specimens are missing their upper spire. Growth lines are hard to discern, usually visible only on a single whorl (typically the penultimate whorl), and were rarely preserved on the base of the last whorl. Some of the specimens appear to have a wider pleural angle (23°) than normal, but these particular specimens have been crushed. A few of the specimens (five percent) have naticid boreholes, and a few other specimens are en- crusted, in part, by bryozoans. Rare spe cimens are coated by calcareous algae. All previous workers assigned Mesalia clarki to various genera without knowledge of the shape of the aperture. Our cleaning of representative specimens of Dickerson’s species revealed it to have a short, shallow effuse spout (Figures 24-25) and bicostate sculpture on the juvenile whorls (F igure 30). There is considerable variation in the sculpture and the shape of the whorls on M. clarki. Some R. L. Squires and L. R. Saul, 2007 Page 11 28 be 20 Figures 24-32. Mesalia clarki (Dickerson, 1914). Specimens coated with ammonium chloride. All from LACMIP loc. 10508. 24. Hypotype LACMIP 13405, apertural view, height 21.5 mm, diameter, 19.6 mm. 25. Hypotype LACMIP 13406, slightly oblique apertural view, height 23.1 mm, diameter 10.8 mm. 26-27. Hypotype LACMIP 13407, height 22.6 mm, diameter 9.4 mm. 26, Abapertural view. 27. Oblique apertural view. 28. Hypotype LACMIP 13408, apertural view, height 23.2 mm, diameter 9.2 mm. 29. Hypotype LACMIP 13409, abapertural view, height 21.5 mm, diameter 11 mm. 30. _Hypotype LACMIP 13410, abapertural view, height 23.6 mm, diameter 9.7 mm. 31. Hypotype LACMIP 13411, base, diameter 9.5 mm. 32. Hypotype LACMIP 13408, base of same specimen shown in Figure 28, diameter 8.9 mm. specimens have nearly uniform sculpture and flattish Zinsmeister (1983: pl. 2, fig. 14), Zinsmeister and whorls (Figure 24), others have carinate whorls (Figure Paredes-Mejia (1958: table 1), ), and Paredes- Mejia (1989: 26). whereas others have uniform sculpture with convex table 3) reported M. clarki from the Santa Susana For- whorls (Figure 29). mation in the Simi Hills, southern California. These re- Page 12 THE NAUTILUS, Vol. 121, No. 1 ports, however, were based on the misidentification of a specimen of Mesalia martinezensis that happens to lack a strong anterior carina on the otherwise convex whorls. Mesalia clarki resembles Motyris aralica (Michailovski, 1912; Wenz, 1939: 652, fig. 1856) from upper Eocene rocks in the Aral Sea region. Motyris Eames, 1952, was formerly known as Tomyris Michailovski, 1912. See Marwick (1957: 162-163) for more taxonomic information pane Motyris. Me salia clarki differs from M. aralica by not having tabulate whorls with strongly indented sutures. The full aperture of M. aralica is enlnowel, and details about its apical whorl development are wanting. The only other species of Motyris that we are aware of is gee ‘is pseudoaralica Eames (1952: 30-31, pl. 1, fig. 15; pl. 2, figs. 58a, b) from Pakistan, but its aperture is unknown. We believe that when the great variability of Mesalia is taken into ac- count, Motyri ‘is will prove to be congeneric. ACKNOWLEDGMENTS Earl Brabb (U. S. Geological Survey, Menlo Park) pro- vided very useful information regar ding the stratigraphy of the beds in the vicinity of the type iscality of Me salia clarki. Lindsey T. Groves (LACM, Malacology Section) kindly provided key literature dealing with the ecology of modem Mesalia. The manuscript benehied from the re- views by Warren D. Allmon (Paleontological Research In- stitute, Ithaca, New York) and Steffen Kiel (University of Leeds, England and Department of Paleobiology, Smith- sonian Institution’s National Museum of Natural History). Steffen Kiel also provided us with an important hard-to- find reference and very useful stratigraphic information. LITERATURE CITED Abbass, H. L. 1963. A monograph on the Egyptian Cretaceous ber United Arab Republic, Ministry of Industry, Geological Survey and Mineral Research Department, Ge ological Museum, Palaeontological Series Monograph 146 pp. ee P.L.. D. P. Smith, W. V. Sliter, and L. R. Saul. 1995. Paleogeography of three Paleocene eens in Baja California, Mexico. In: A. E. Fritsche (ed.) Cenozoic Pa- oe of the Western United States—II. Pacific Section (Society for Sedimentary Geology), Book 75: 1-S. Adams, A. and L. A. Reeve. 1848-1850. Mollusca In: A. Adams (ed.) The Zoology of the Voyage of H. S. Samarang, under the Command of Captain Sir E e - Belcher ... 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Bulletin of the American Museum of Natural History 3(2): 381-441. Page 16 THE NAUTILUS, Vol. 121, No. 1 Wiedey, L. W. 1928. Notes on the Vaqueros and Temblor formations of the California Mioce ne with descriptions of new species. Transactions of the San Diego Society of Natural History 5(10): 95-182. Woods, H. 1922. Mollusca from the Eocene and Miocene de- posits of Peru. In: T. O. Bosworth (ed.) Geology of the Tertiary and Quaternary periods in the north- west part of Peru. MacMillan and Company, London, pp. 51-139. Zinsmeister, W. J. 1974. Paleocene biostratigraphy of the Simi Hills, Ventura County, California. University of California, Riverside, unpub. Ph. D. dissertation, xii + 236 pp. Zinsmeister, W, J. 1983. Late Paleocene (“Martinez Provincial Stage”) mollusean fauna from the Simi Hills, Ventura eae California. In: R. L. Squires and M. V. Filewicz (eds.) Cenozoic Geology of the Simi Valley Area, Southern Califomia. Pacific Section, SEPM Fall Field Trip Volume and Guidebook, Book 35, pp. 61-70. Zinsmeister, W. J. and L. M. Paredes-Mejia. 1955. Paleocene biogeography of the west coast of North America: A look at the molluscan fauna from Sepultura Formation, Mesa San Carlos, Baja California Norte. In: M. V. Filewicz and R. L. Squires (eds.) Paleogene Stratigraphy, West Coast of North America. Pacific Section, SEPM West Coast Paleo- gene Symposium, V olume 58, pp. 9-22. APPENDIX LOCALITIES CITED Localities are LACMIP, unless otherwise noted. All quadrangle maps are U. S. Geological Survey maps. 7060. Elevation 1427 ft., on ridge between Temesal and Santa Ynez canyons at edge of fire road on top of ridge, Topanga Canyon ‘Quadrangl e (7.5 minute, 1952, photorevise od 1981), Los Angeles County, southern California. Pale- ocene. Santa Susana Formation. Coll.: H. D. B. Wilson, June 1, 1941. 10508. North slope of Trailer Canyon near top of ridge be- tween Quarry and Trailer canyons at approximately 1325 ft. elevation and just west of saddle, just below coralline- algal beds in limy siltstone west of small f fault, road cut north side of unpaved road 5600 ft. north of San Vicente y Santa Monica Grant boundary, 10,400 ft. east of Los Angeles City boundary, Topanga Quadrangle (7.5 minute, 1952, photor vised 1981), east of Santa Ynez ¢ Janyon, Pali- sades Highlands, Santa Monica Mountains, Los Angeles County, southern California. Lower upper Paleocene (lower Thanetian). Santa Susana Formation. Coll.; ¢ Strathearn and others, fall, 1982. 11717. Float at about 1600 ft. elevation in bottom of south- flowing gully joining Quarry Canyon at about 1410 ft. =“ evation: 1500 ft. SW of hill 2036, Topi nga Quadrangle ( minute, 1952, photorevised 1981), Los Angeles C a southern C ali tonmie Paleocene. Santa Susana Formation. Coll J.M Alderson, November 11, 1980. 21581. Black nodular shale and conglomerate on road 1.1 mi. east from Cienaga Camp at Fish Canyon forks toward Warm m Springs summit: on northwest side of ravine: north side East Fork Fish Canyon, T. 6 N, R. 16 W, approximately at. ft. north, 750 ft. east of bench mark 2205, Warm Springs Mountain Quadrangle (7.5 minute, 1958, photo- revised 1974), Los Angeles County, southern California. Pa- leocene ( (upper Danian). San Francisquito Formation. Coll: R. W. Webb and E. H. Quayle, jane 1941, 21607. South 1/2, SE 1/4, section 25, T. 2 N, R. 18 W, Cala- basas Quadrangle (7.5 minute, 1952, photorevised 1967), Ventura County, southern California. Lower upper Pale- ocene (lower Thanetian). Santa Susana Formation. Coll: Kinney and Sherman, date unknown. 21554. Reworked fossil boulders in conglomerate, in second spur canyon off Poison Oak Canyon east of Las Llajas Canyon, 2800 ft. up canyon (north) from Poison Oak Can- yon; on west slope 25 ft. above bottom of canyon, Santa Susana Quadrangle (7.5 minute, 1951, photorevised 1969), north side Simi Valley, Ventura County, southern Califor- nia. Paleocene. Santa Susana Formation, Coll.; P. L. Gold- man, date unknown. 22330, Beds cropping out on nose of spur on west side of Meier Canyon, approximately 600 ft. north of second “n” in “Meier Canyon,” Santa Susana Quadrangle (7.5 minute, 1951, photorevised 1969), south side of Simi Valley, Simi Hills, Ventura County, southern California. Lower middle Paleocene (Selandian) = Turritella peninsularis Zone. Santa Susana Formation, “Martinez marine member.” Coll.: W. P. Popenoe, April 3, 1946. 22557. Sandstone bed below small waterfall [dry] west of road going south through Barclay Ranch, 10,162 ft. south and 5660 ft. wet of junction of Souther Pacific railroad and Los Angeles Ave. about 0.25 mi. east of Santa Susana, Santa Susana Quadrangle (7.5 minute, 1951, photorevised 1969), Simi Hills, Ventura County, southern California. Paleocene. Santa Susana Formation. Coll: M. Murphy, spring, 1950. 22698. On first large ridge; trending southwest to west of ridge trending south of hill 2150. Bearing from the nioihowest corner of the Calabasas Quadrangle is S14°E; distance 12.210 ft., Calabasas Quadrangle (7.5 minute, 1952, pho- torevised 1967), Simi Hills, Ventura County, southern California. Paleocene. Santa Susana Formation. Coll.: J. H. Fantozzi, June 1, 1951, 25281. Sandstone at elevation of 1000 ft., about 400 ft. south aiid LO0O0 ft. west of northeast corner of section 5, T. 1S, R. 17 W, Malibu Beach Quadrangle (7.5 minute, 1950, photorevised 1967), on west side of northern tributary to Stokes C anyon, western Santa Monica Mountains, Los ‘Ane geles County, southerm California. Reworked Paleocene (Selandian) fossils in middle Miocene Calabasas Forma- tion, Stokes Canyon Breccia Member. Coll.: J. Stark and T. Susuki family, May 5, 1965. 26897. Gully west side of Temesal Canyon opposite 2nd ‘e’ of Temesal at about 1475 ft. elevation; approximately 1082 m (3550 ft.) south; 533m (1750 ft.) east of hill 22036; San Vincente and Santa Monica Grant, Topanga Quadrangle (7.5 minute, 1952, photorevised 1967), Santa Monica Mountains, Los Angeles County, southern California. Middle upper Paleocene (middle Thanetian). Santa Su- sana Formation, Coll: ]. M. Alderson, March 9, 1980. UCMP. 1540. Elevation 1000 ft., 1 mi. south of Stewartville (site), northeast corner of NW 1/4, section 15, T. 1 N, R. 1 E, Antioch South Quadrangle (7.5 mimute, 1973, pho- torevised), 300 ft. south of basal Tejon conglomerate and 600 ft. north of Chico-Martinez contact, northeast side of Mount Diablo, Contra Costa County, northern California. Upper middle Paleocene (Selandian) = Turritella infra- granulata pac hecoensis ae “Martinez” Formation, lower member. Coll: | ). Dickerson, circa 1912. THE NAUTILUS 121(1):17-28, 2007 Page 17 Vertigo malleata, a new extreme alcifuge land snail (Gastropoda: Vertiginidae) from the Atlantic and Gulf coastal plains of the USA Brian F. Coles Mollusca Section, Department of Biodiversity National Museum of Wales, Cathays Park Cardiff, CF10 3NP , GREAT BRITAIN pristiloma@hotmail.com Jeffrey C. Nekola Department of Biology University of New Me xIiCO Albuque rque, NM $7131 USA jnekola@unm.edu ABSTRACT Vertigo malleata new species is an extreme calcifuge land snail widely distributed in the Atlantic and Gulf costal plains of the eastern USA. This species appears to have gone undetected because of its small size and restriction to low pH = sites— —— bogs, Atlantic white cedar (Chamaecyparis as S L.) BSP) swamps, pocosins, and pine woodlands—which, has been assumed, harbor little or no molluscan diversity. Ver r tigo malleata is distinguished from other members of the genus by the strongly pustulose surface of the body whorl, which gives the shell a malleated appearance at low to moderate mé vonifi- cation. While the major apertural lamellae/folds (parie tal, col- umellar, and palatal) of this species are typical for Vertigo, the strongly pustulose shell sculpture, occurrence of an infrapari- etal lamella, and frequent development of subcolumellar and basal lamellae in the absence of an angular lamella appear unique. Although V. malleata is an abundant snail within its range, the common use of short-return fire regimens to manage forests of the eastern USA appears to be artificially limiting its distribution to wet, less frequently bumed sites. Additional key words: Bothriopupa, Nesopupa, biogeography, fire ecology, community ecology, eastern North America INTRODUCTION Acidic and lime-poor habitats have long been thought to support depauperate molluscan community abundance and richness (Boycott, 1934; Baker, 1939; Kerey and Cameron. 1979; Burch and Pearce, 1990). Consec quently, little molluscan survey work has been attempted in acidic sites even though they can re present a substantial frac- tion of the landscape. However, such areas should not be ignored for terrestrial gastropod biodiversity because base-poor habitats can be as speciose as base-rich habi- tats on a per-individual basis (Schilthuizen et al., 2003; Corresponding author: Jeff Nekola email: jnekola@unm.edu Pokryszko and Cameron, 2005), and because some spe- cies like the European Vertigo ronnebyensis (Wester- lund, 1871) and Zonitoides excavatus (Alder, 1830) are restricted to or more frequent in bea sites (Kemey and Cameron, 1979), During land snail studies in eastern North America (Nekola, 2002a; Nekola and Coles, 2004; Coles and Nekola, unpublished data) we found that acidic habitats often supported substantial populations of land snail taxa that have been little reported since their original descrip- tions; e.g., Vertigo cristata (Sterki, 1919), Vertigo nylan- deri Sterki, 1909, Vertigo alabamensis Clapp, 1915, and Vertigo perryi Sterki, 1905. In tact, V. alabamensis and V. perryi were each previously known from only two sites worldwide (Pilsbry, 1948: Hubricht, 1985). In the course of these acid-habitat surv eys, we examined Saco Heath, an undisturbed domed ombotrophic Sphagnum bog in the Atlantic coastal plain of York County, Maine. At this site we located a species of the genus Vertigo that was strikingly different from all previously inoue taxa. This form was subsequently found to represent the most com- mon land snail of highly acid, mesic to wet habitats of the Atlantic and Gulf coastal plains of the eastern USA. Here, we describe this taxon as Vertigo malleata, new species, document its biogeography and ecology, and briefly discuss its relevant conservation issues. MATERIALS AND METHODS Site Selection: Approxiini itely 130 sites were surveyed along the Atlantic and Gulf coastal plains of the eastern USA from Maine to western gba i, including penin- sular Florida south to Gainesville. These sites encom- passed the entire soil base-status and moisture gradient of the region and covered a total geographic extent of 2400 kin. Thirty sites represented base-neutral to base- rich habitats (i.e., forests, and limestone outcrops), while the remaining were base-poor (2.e., pine barrens, pine-wiregrass Sda- calcareous wetlands, marl banks, rich Page 18 THE NAUTILUS, Vol. 121, No. 1 vanna, heaths, Atlantic white cedar swamps, bay forest, Sphagnum bogs, and pocosins). Field Methods: Latitude and longitude of each site was determined using a hand-held GPS. Terrestrial gas- tropod faunas were documented from a representative 100-1000 m2 area within each site by hand collection of larger taxa and litter sampling for smaller taxa. Litter sapling was used as the primary method of collection because it provides the most complete assessment of site faunas (Oggier et al., 1995; Cameron and Pokryszko, 2005). As suggested by Emberton et al. (1996), collec- tions were medé at places of high micro-mollusk density such as loosely compacted leaf litter lying on top of highly compacted damp soil or humus. This loose litter was removed by hand and aggressively sieved in the field using a shallow sieve of 2 mm mesh nesting loosely inside a sieve of 0.6 mm mesh. The procedure consisted of throwing handfuls of litter onto the coarser mesh accom- panied by vigorous shaking, tapping, or other agitation. The process was continued for 15-60 minutes, a time interval that yielded 50-500 ml of fine material (0.6—2.0 mm). In general, sites were sampled in parallel (but in- dependently) by each of the authors, although several sites were sampled by only one worker (see Table 1) Laboratory Procedures: Samples were slowly and completely ‘dried at room te mperature and then passed through an ASTME #30 sieve (0.6 mm mesh) with frac- tions be sing hand-picked against a neutral background. All shells, shell fragments, and slug plates were removed, and all identif fable shells from each site were assigned to species using the authors’ reference collections and vari- ous museum collections (see below). The total numbers of shells per species per site were recorded, as were the number of unidentified immature individuals. Comparisons: The new species was compared with specimens of all eastern North American and western Eurasian species of Vertigo, and to representative taxa in the related genera Nearctula of western North America, Nesopupa of the Old-World tropics, and Bothriopupa of the neotropics. Comparative material consisted of the authors’ extensive reference collections, the collections of the gees Museum of Natural History, Gainsville, ): the Field Museum of Natural History, Chi- cago, . (FMNH), the Carnegie Museum of Natural History, Pittsburgh, PA (CM); the Wales—Zoology, Cardiff, U K (NMW.Z): the Natural His- tory Museum, London, UK, and the Queensland Mu- seum, Brisbane, Australia. Additional comparisons were made with material presented by Pilsbry (1920; 1948). Imaging: ing a digital camera attached to a stereomicroscope. Ap- proximately 12 separate 1388 x 1040 pixel images were made of each specimen with the image focal lengths Shells were imaged at 40x magnification us- positioned at 120%m increments from the front to back of the shell. CombineZ5 freeware (ht ttp:/Avww. hadleyweb p.blueyonder.co.uk/CZ5/ combinez5.htm) was used see ese National Museum of to assemble a final image from the well-focused parts of each separate image, The body whorl surface of the new taxon was also imaged at 150x with 60 images positioned at 5 jum focal length increments and assembled into a single image using CombineZ5. These separate images were imported into Adobe Photoshop, where brightness and contrast were optimized and the background made uniformly black. These images were then compiled into a single plate. Community Ecology: — Analysis of co-occurring terres- trial gastropod species and abundance was determined using data for sites sampled by the second author (ie., all . s with accession numbers prefixed by JON in Table ). These analyses were performed for the whole dataset of 49 discrete sites and also by geographic sub-region— New England (Maine, Massachusetts), New Jersey, the North and South Carolina coastal plain, and the Gulf coastal plain—to allow for documentation of composi- tional gradients across the range of the new species. The physical habitat and plant community from each site was also noted. Nomenclature: Taxonomic nomenclature follows that of Turgeon et al. (1998) with updates from Nekola (2004). Apertural lamellae and fold nomenclature follows that of Pilsbry (1945: 869, fig. 469), i.e., parietal “teeth” are referred to as “folds” and all other “teeth” are termed “lamellae”, whatever their form. SYSTEMATICS Class Gastropoda Subclass Pulmonata Order Stylommatophor l Family Vertiginidae Genus Vertigo Miiller, 1773 Vertigo malleata new species (Figures 1-15, 20, Tables 1-2) Diagnosis: Minute; shell ovoid, similar in size and shape to Vertigo ventricosa (Morse, 1565) but. distin- cuished by malleated appearance of the body whorl at low to moderate (10-40x) magnification; upper whorls fine ‘ly rib-striate, minutely decussated by spiral lines; ap- erture with parietal and columellar lamellae, a small in- fraparietal lamella (occasionally absent), and two palatal folds; one or more subcolumellar-basal lamellae usually present; angular lamella absent. Description: Shell 1.5—2.1 mm tall x 1.25-1.4 mm wide (holotype 1.98 x 1.36 mm), ovoid to ovoid-conical, inflated, approximately 44.5 whorls, with deep suture; translucent, olive-ye low to brown in color; body whorl approximately 66% of total he ight. Protoconch and neanic whorls minutely papillose pre fine spiral stria- tion; subsequent whorls finely rib-striate: striae most dis- tinct on penultimate whorl where they are minutely de- cussated by fine spiral lines; on body whorl the se ulpture degenerates into an irregularly pustulose surface (Figure B. F. Coles and J. C. Nekola, 2007 Page 19 4) which at low to moderate (10-40x) magnification takes on a malleated appearance as it appears hammered with small depressions; behind the aperture the sculp- ture takes the form of coarse, irregular rib-striae (Figures 2. 11). Aperture rounded, approximately 40% of shell height; lip reflexed but not thickened, peristome usually dark blackish-olive: sinulus moderate-weak: basally the aperture abruptly inflates to form a rounded swelling, but not a distinct crest (Figures 2, 8). Umbilicus closed (Figure 3). Aperture typically with six lamellae and folds (Figures 1, 5, 7, 9, 12-15): a strong, slightly sinuous pa- rietal lamella (Figures 1,5, 9, 15); a shelf-like columellar lamella that spirals internally around the columella for approximately one whorl; two palatal folds of approxi- mately equal length that extend approximately 0.2 whorls into body whorl, the lower slightly more immersed than the upper (Figures 1, 5-7, 10, 15), both highest at mid- length (Figures 1, 5, 6, 15); a nodular infraparietal lamella usually present (Figures 1,5, 9, 12-15), occasion- ally vestigial or absent (Figures 6, 10); angular lamella absent: presence of a nodular subcolumellar lamella and nodular subcolumellar-basal lamella variable (Figures 1, 6, 7, 9, 10). Apertural ends of the palatal folds coincide with abrupt inflation of basal aperture (Figures 7-8), in consequence appearing to be raised on a weak crest when viewed within the aperture but not associated with any internal shell thickening; externally shell only slightly impressed over palatal folds. Body of animal grey vat several organs of a brown or cream color visible through the upper whorls of shell. All dissected individuals bias e proven to be aphallic (Beata Pokryszko, personal com- munication), hence the genitalic anatomy is unknown. Holotype (Figures 1-4): NMW.Z.2005.011.03830, USA North Carolina, Pender County, Holly Shelter Game Land, Brian Coles, 1 April 2003. peas (Figures 5-15): || NM\W.Z.2005.011.03531- 03839, figured material, see Figure legends for details; NMW.Z.2005.011.02118-02120, approximately 5100 in- dividuals (split into three approximately equal lots) from type locality; UF 348143, approximately 700 individuals from type locality; CM 73971, 143 individuals from type locality; NMW Z,.2005.011.02597, 90 specimens, Wells Heath, York County, Maine (43°20'2” N, 70°38'24” W), Brian Coles: NMW.Z.2005.011.02591, 26 specimens, Skunknett Audubon Preserve, Barnstable County, Mas- sachusetts (41°38'59” N, 70°22’31” W): NMW.Z.2005. 011.02585, 170 specimens, Peterson Swamp Wildlife Management Area, Plymouth County, Massachusetts (42°0'37" N, 70°49'4” W), Brian Coles: NMW.Z.2005. 011.02514, 122 specimens, Stafford Forge Wildlife Man- agement Area, Ocean County, New Jersey (39°42'44" N, 74°22'10" W), Brian Coles: NMW.Z.2005.011.02197, 250 specimens, Lewis Ocean Bay Preserve, Horry County, South Carolina (33°47'16" N., 78°50'56" W.), Brian Coles; NMW.Z.2005.011.03035, 42 specimens, Collins Bay, Ware County, Georgia (31°5'12” N., §2°36'56" W.), Brian Coles: NMW.Z.2005.011.03065, 107 specimens, Wilma Station, Liberty County, Florida (30°9'34" N., $4°57'39" W.), Brian Coles; NMW.Z.2005. 011.03079, 162 specimens, Pond Creek, Conecuh Na- tional Forest, Covington County, Alabama (31°6'12" N., 86°32'3" W.), Brian Coles. Type Locality: Holly Shelter Game Land (34°31'57" N, 77°44'41” W), Pender County, North Carolina, USA; ae dense scrub of mesic bay/pine forest at pocosin margin, individuals sieved from deep bracken fern and pine needle litter, collected by Brian Coles, 1 April 2003. Other Material (Table 1): Sixty additional lots col- lected by Brian Coles are deposited i in the Coles Collec- tion of the National Museum of Wales. Fifty three lots representing 3133 individuals collected by Jeff Nekola are deposited in the Nekola collection (JCN). Etymology: The specific name malleata refers to the hammered appearance of the body whorl at low to mod- erate magnification. Variation: Vertigo malleata was rather constant in general appearance along its 2400 km range, although some variation in shape, size, color, sculpture, and de- velopment of the apertural lamellae was observed. Varia- tion in size and shape has been noted above. In addition, the most southern populations (Georgia, Alabama, and Florida) tended to be darker in color and showed the most strongly developed shell sculpture (Figures 9-12). While the parietal lamella, columellar lame: and the palatal folds varied little, the infraparietal lamella varied from strong (Figures 1, 5, 19) to weak (Figure 9) to occasionally warty (Figures 6,10). The eae and nodular basal lamella although usually distinct (Fig- ures 1, 5, 7, 13) were also occ: isionally absent ( Figure 5). Multiple subcolumellar-basal lamellae of eS place- ment were also noted most frequently in Gulf Coast populations (Figures 9, 10). However, such trends were not distinct enough to support the designation of geo- graphical races, w ith most of this observ ed morphological variation occurring within local regions or populations. Comparison with Other Species of Vertigo and of Related Genera: Vertigo malleata differs from all other Vertigo species by its strongly pustulose body whorl sculpture and possession of an infraparietal and subcolumellar-basal lamellae while lacking an angular lamella. Because of these unusual characteristics, we do not feel assignment of this taxon to a particular subgenus to be prudent at this time. Additional data, possibly based on DNA sequence information, will be required to accurately determine its closest relatives. On casual inspection, Vertigo malleata could be taken for a member of the V. gouldii group (e.g. Vertigo cris- tata; see Pilsbry, 1948: 958, figs. 4, 5, 8; 967, figs. 1-16) because of its shell color, striated upper whorls, and silky luster. Like V. malleata, V. cristata has four prominent lamellae and strong striation on the penultimate whorl (Pilsbry, 1948: 967, figs. 4-5, 973, fig. 520; Nekola, 2001) Page 20 THE NAUTILUS, Vol. 121, No. 1 Table 1. Vertigo malleata: sites, brief habitat descriptions, collection dates, accession numbers, and total number of specimens taken. Site State/County # Site: Habitat! Coordinates Date Accession Nufmber Specimens Alabama Covington 1 Pond Creek seep (Conecuh — 31°06'12" N, May 5 2005 NMW.Z.2005.011.03079 162 NF); Tlex-Smilax-bay 86°32'03" W JON 12365 LU scrub on seep margin 2 Moccasin Branch (Conecuh 31°06'42" NJ May 5 2005 JCN 12371 3 NF); old pine-bay-heath 86°35'53" W forest 3 Bear Bay (Conecuh NF): 31°6'29" N, May 5 2005 = NMW.Z.2005.011.03068 ] heath-dominated 86°38'54" W scrub on wetland margin Mobile 4 Grand Bay Forever Wild 30°25'07" N, May 1 2005) NMW.Z.2005.011.03019 3 Preserve; wet bay and $8°19'35" W mixed forest Florida Columbia 5 Impassable bay (Osceola 30°23'31" N, Jan § 2005 NMW.Z.2005.011.02849 740 NF WMA); wet holly-bay — $2°30'05" W May 2 2005 JCN 12280 71 scrub 6 Osceola National Forest 30°22'30" N, May 2 2005) NMW.Z.2005.011.03026 213 WMA; wet Pinus- 82°32'04" W JON 12285 75 Lyonia-Vaccinium savanna 7 ~~ Osceola National Forest 30°22'39" N, Jan § 2005 NMW.Z.2005.01 1.02845 38 WMA: wet Acer- 82°31'42" W May 2 2005 | NMW.Z.2005.011.03024 23 Taxodium-Lyonia forest May 2 2005. JCN 12283 6 Leon S Wolf Trap Bay 30°22'04" N, Jan 7 2005 NMW.Z.2005.011.02513 ~100 (Apal: achicola NF); tall S4°34'11" Wo May 4 2005) NMW.Z.2005.011.03054 132 pine-holly-bay forest May 4 2005 JCN 12324 37 9 Wolf T rap Bay 30°21'46" N Jan 7 2005 NMW.Z..2005.01 1.02516 44 (Apalachicola NF): $4°34/23" Wo May 4 2005 NMW.Z.2005.011.03050 7 wet-mesic pine-holly- May 4 2005 JCN 12321 1] heath forest 10 Otter camp (Apalachicola 30°20'20" N, Jan 7 2005 NMW.Z.2005.011.02520 ~50 NF); regenerating mesic S4°36'41" Wo May 4 2005) NMW.Z.2005.011.03056 4 pine-holly heath Liberty 1] Wilma Station: mesic, old 30° °09'34" N, May 4 2005 = NMW.Z.2005.011.03065 127 pine-magnolia-bay forest °57'39" W JON 12344 30 12 Juniper Creek Islands 30°03'15" N, May 4 2005) NMW.Z.2005.011.03062 65 (Apalachicola NF); old 4°45'40" W JCN 12337 41 growth pine-holly-bay forest 13° Juniper Creek Islands 30°04'46" N, May 4 2005) NMW.Z.2005,011.03059 5 (Apalachicola NF); S4°45'41" W JON 12333 75 white cedar-pine-holly forest I4 Juniper Creek Islands 80°02'07" N, May 4 2005 NMW.Z.2005,011.03064 ~40 (Apalachicola NF); 84°49'38" W JON 12539 4S pine-red maple-white cedar forest 15 Carr Bridge (Apalachicola 30°07'26" N, May 4 2005. JCN 12340 12 NF); wet-mesic Ilex 84°53/31"” W forest Wakulla 16 South of Otter Camp 30°16'55" N May 4 2005) NMW.Z.2005.011.03057 69 (Apalachicola NF); $4°36'54" W May 4 2005 JCN 12327 53 wet-mesic pine-holly forest 17 W Branch Sopchoppy R 30°15/05" No May 4 2005 NMW.Z.2005.011.03058 5 (Apalachicola NF); $4°37'30" W JON 12328 36 pine cy press-bay-holly forest B. F. Coles and J. C. Nekola, 2007 Page 21 Table 1. Continued Site State/County # Site; Habitat! Coordinates Date Accession Number Specimens Georgia Ware IS Collins Bay; wet holly-wax 31°05'12" N, May 2 2005 NMW.Z.2005.011.03035 42 myrtle-bay forest §2°36'56" W JON 12300 58 19 Dixon State Forest; wet 31°05'36" N, May 3 2005 JON 1230] 4 Pinus-Gordonia forest 82°16'13" W 20 Dixon State Forest; 31°06'49" N, May 3 2005 NMW.Z.2005.01 1.03035 ] wet-mesic Quercus- 82°16'16" W JON 12302 2 Ilex-Gordonia forest Maine York 2) Saco Heath 1 (TNC): sedge 43°32'42" N Oct 14 2002 NMW.Z.2005.01 1.01550 3 and heath litter on 70°28'33" W Aug § 2004 NMW.Z.2005.01 1.02567 l4 Sphagnum bog NMW.Z.2005.011.02577 52 JON 12092 10 JON 12099 88 JON 12101 3 Oct O1 2004 NMW.Z.2005.011.02614 55 NMW.Z.2005.011.02616 40 22 Saco Heath 2 (TNC); low 43°32'50" N, Aug OS 2004. NMW.Z.2005.011.02571 45 forest with Carex 70°27'32” W JEN 12095 59 groundcover ; 23. Wells Heath (TNC); under 43°20'02" N, Oct 01 2004. NMW.Z.2005.011.02597 90 heath scrub on 70°38'24" W Sphagnum bog Massachusetts Barnstable 24 Skunknett Audubon 41°38'59" N, Aug 13 2004. NMW.Z.2005.011.02591 26 Preserve 2; Myrica- 70°22'31"” W JCN 12180 22 Chamaecyparis bog margin Bristol 25 Noquochoke WMA: 41°39'35" N, Aug 12 2004. NMW.Z.2005.011.02592 2) Chamaecyparis-Cryilla T1°O1'07" W JON 12168 17 swamp forest 26 Noquochoke WMA: 41°39'39" N, Aug 12 2004 NMW.Z.2005.01 1.02590 4 leatherleaf island in acid 71°01 12" W JCN 12164 12 sedge fen 27 Pine Swamp Brook; 41°55'57" N, Aug 10 2004. NMW.Z.2005,011.02586 9 leatherleaf fringe of open 71°03'49" W JCN 12149 5 acid bog Plymouth 28 Peterson Swamp WMA; 42°00'37" N, Aug 10 2004. NMW.Z.2005.011.02585 ~170 Chamaecyparis-Acer 70°49'04" W JON 12145 137 rubrum forest Worcester 29. Tom Bog; Sphagnum bog 42°30'46" N, Oct 05 2004. NMW.Z.2005.011.02760 with serub 72°12'43”" W NMW.Z.2005.011.02761 ~170 New Jersey Atlantic 30 Park Road (Wharton SF): 39°42'58" N, May 22 2004 = NMW.Z.2005,011.02516 18 moist Ilex-Gaylussacia- 74°44'10" W JON 12050 54 Kalmia scrub Burlington 31 Swan Bay WMA; low 39°35'13" N May 20 2004 =NMW.Z.2005.011.02479 107 Nyssa-llex-. Acer rubrum 74°30'50" W JEN 11983 103 forest 32 Lebanon State Forest; open — 39°52’28" N, — May 19 2004. =NMW.Z.2005.011.02469 2] heath-Smilax scrub 74°30'57" W May 21 2004 JCN 12026 l4 33. Roberts Brook; low Nyssa- 39°47'07" N, May 19 2004 =NMW.Z.2005.011.02466 20) Chamaecyparis-heath 74°39/26" Ws May 21 2004. JCN 11989 5 forest 34 Brendan T Byrne State 39°53'07" N, May 22 2004. NMW.Z.2005.011.02499 | Forest: dry 74°30'22" W Chamaecyparis-bog Camden 35 2 miles WSW of Delette: 39°46'32"” N, May 19 2004 = NMW.Z.2005.011.0246] 6 moist bank with pine, 74°48'21" W May 21 2004 = NMW.Z.2005.011.02454 28 oak, wax myrtle 8 JON 11995 THE NAUTILUS, Vol. 121, No. 1 Table 1. Continued Site State/County # Site; Habitat! Coordinates Date Accession Number Specimens Gloucester 36 Winslow WMA: heath scrub 39°37'08" N, May 23 2004 = NMW.Z.2005.011.02518 12 in abandoned blueberry 74°53'43" W JON 12054 14 field Ocean 37 Colliers Mill WMA: acid 40°05'35" N, May 22 2004. NMW.Z.2005.011.02508 ~150 bog with Ilex, 74°25'58" W JEN 12036 73 Chamaedaphne, Aronia 38 Stafford Forge WMA; moist —39°42'44" N. May 22 2004 = NMW.Z.2005.011.02514 122 Kalmia-Pinus forest 74°22'10" W JON 12045 54 39°53'34" N May 22 2004 = NMW.Z.2005.011.02511 3 74°19'58" W JEN 12039 15 North Carolina Bladen 39 sas Mill Bay (Bladen eas N Jun 02 2003 | NMW.Z.2005.011.02204 ~100 uakes SF); pocosin with 31/33” W JEN10613 142 ae yparis Brunswick 40 Green Swamp (TNC); 34°06'14" N. fun O01 2003) NMW.Z.2005.011.02193 50 Chamaecyparis-bay 78°18'35" W JON 10615 94 forest 41 Green Swamp (TNC); 34°05'42" N, Jun O01 2003) | NMW.Z.2005.011.02194 20 medium pocosin 78°17'48" W JON 10617 19 42 Prospect Ridge; mature 34°03'48" N, Jun OL 2003 NMW.Z.2005.011.02196 25 pine-bay forest 78°20'52" W JCN 10622 2 Carteret 43 Millis Road (Croatan NF): 34°46'16" N, Feb 24 2003) NMW.Z.2005.011.02128 82 wet pocosin with 76°58'39" W JCN 10624 leatherleaf Craven 44. Sheep Ridge (Croatan NF); 34°56'07" N, Feb 24 2003) NMW.Z.2005.011.02132 ~600 medium pocosin 77°04' 14" W NMW.Z.2005.011.02130 30 JON 10693 48] JON 10708 10 45 Catfish Lake South 34°55'39" N. Feb 24 2003. =NMW.Z.2005.011.02126 ~400 (Croatan NF); low, wet 77°05'05" W JON 10675 257 pocosin 46 Catfish Lake South 34°55'10" N, Feb 24 2003 JCN 10668 l (Croatan NF): roadside 77°05'24" W ditch in medium pocosin 47 Neusiok Trail North 34°54'03" N, Jun 01 2003) NMW.Z.2005.011.02190 20 (Croatan NF); wet-mesic 76°49'06" W JCN 10686 pine-bay forest ones 48 Catfish Lake Wildemess 34°55'07" N, Feb 24 2003) NMW.Z.2005.011.02125 40 (Croatan NF); mature 77°10'43" W JCN 10713 64 bay-pine forest Moore 49 Pinebluff; bay forest in 35°06'14" N, Jun 03 2003 JON 10746 is) gulley along US 1 79°28'28" W Pamlico 50 Goose Creek Game Land: 85°15'14" N, May 31 2003 | NMW.Z.2005.011.021S8 10 pine straw under scrub 76°35'52" W Pender 51 Holly Shelter game land; 8157" N, Apr O1 2003) NMW.Z.2005.011.03830 Holotype edge of mature mesic 76: 4’ 11" W NMW.Z.2005.011.02119 ~1700° bay/pine forest NMW.Z.2005.011.02118 ~1700° NMW.Z.2005.011.02120 ~1700° CM73971 143° UF348143 ~700° 52 Holly Shelter game land; 34°32'57" N, Apr O1 2003) | NMW.Z.2005.011.02428 ~600 dense pocosin scrub 77°46'54" W 53 Holly Shelter game land; 34°33'06" N, Apr O01 2003 NMW.Z.2005.011.02117 115 dense bay scrub 77°47'37" W 54 Lanier Quarry (TNC); 34°37'49” N, Jun OL 2003 NMW.Z.2005.011.02192 ~SO Shrubs bordering 77°40'27" W JON 10753 64 pime-wiregrass Savanna rrell 55 Pocosin Lakes NWR; low 35°42'30" N, Apr 03 2003) NMW.Z.2005.011.02122 ~900 pocosin 76/11" W NMW.Z.2005.01 1.02123 ~9O0 May 31 2003 JCN 10824 92 B. F. Coles and J. C. Nekola, 2007 Table 1. Continued State/County # Site; Habitat! Coordinates Date Accession Number Specimens 56 Pocosin Lakes NWR: maple-oak-pine woodland 35°40'19" N, 76°12'16" W Apr 03 2003 NMW.Z.2005.011.02121 50 57. Frying Pan Landing 35°48'03" N, Apr 03 2003 | NMW.Z.2005.011.02174 | (Pocosin Lake NWR); 76°06'00" Wo May 31 2003 JEN 10828 l pine pocosin ; South Carolina Horry 58 Lewis Ocean Bay Preserve; 33° ee N, Jun 02 2003 NMW.Z.2005.011.02201 ~200 medium pocosin on 78°50'36" Wo JON 10955 300 roadside 59 — Lewis Ocean Bay Preserve; 33°47'16" N, Jun 02 2003 NMW.Z.2005.01 1.02197 ~250 mesic pine-bay forest 78°50'56" W JON 10960 123 60 Lewis Ocean Bay Preserve; 33°47'33" N, Jun 02 2003 NMW.Z.2005.011.02200 95 mesic longleaf pine forest — 78°51'02” Ww JCN 10964 26 ' Abbreviations used are: NF National Forest, NWR National Wildlife Refuge, SF State Forest, TNC The Nature Conservancy, WMA wildlife management area. However, V. cristata has striate (not pustulose) sculpture on the body whorl, has a weak crest (rather than a basal inflation), lacks an infi raparietal lamellae, and has a nodu- lar (not shelf-like) columellar lamella. These two species were found co-occurring in several New England loca- tions (Table 1, sites 21, 92, 23. and 29), where they could readily be distinguished under low m: agnification. Vertigo mulled also resembles Ve rtigo ventricosa (Morse, 1865) and Vertigo perryi Sterki, 1905 with re- spect to the ovoid shape, large aperture vs. shell height ratio, reflected lip (Pilsbry, 1948: 958. figs. 1-3, 7), and basal apertural inflation (Coles and Nekola mpnen data); V. perryi also has a dark colored peristome (Pils- bry, 1948: 968). However, these species cannot easily be confused because V. ventricosa and V. perryi have glossy shells with only weakly developed striae, lack an Ear parietal lamella, and have a peg-like columellar lamella. Although not previously reported in the genus Vertigo, the pustulose sculpture ‘of the body whorl in Vertigo malleata is not unique to this species; Vertigo iE ee sis and Vertigo conecuhensis (Pilsbry, 1948: 949, fig. 510, 9, 12-14: 950, fig. 511) of Ree ee North America also weakly exhibit this trait (Figures 18, 19). At low magnification, the sculpture of V. malleata also some- what resembles the pitted or granular surface of mem- bers of the Nesopupinae. However, members of the Nesopupinae commonly have an angular lamella (i.e., Nesopupa, Sterkia), while none are aown to have an intraparietal lamella. Supertficially, V. malleata also ap- pears similar to the neotropical genus Bothriopupa (Pils- bry, 1948: 1011, fig. 539). However, with respect to shape, color, nature of the surface sculpture and configu- ration of the major apertural lamellae and folds, V. mal- leata much more closely resembles other members of the genus Vertigo (Figures 16, 17). Geographic Distribution: Vertigo malleata occurs from southern Maine to southeastern Georgia along the Atlantic coastal plain to the west side of Mobile Bay along the Gulf coastal plain, apparently excluding pen- =acuiler Florida (Table 1, Figure 20), This distribution includes a number of regions of particular ecological in- terest and conservation concern, e.g. es Pine Barrens of New Jersey (sites 30, 32-35, 37-38), the sandhills and pocosins of the Nou Carolina ie South Carolina coastal plain (sites 39-48, 51-60), the Okefenokee Swamp of southeastern Georgia (sites 19, 20), and the Appalachicola sand plain of western Florida ( (site s 8-17). It seems likely that its distribution extends into the Gulf coastal plain of Mississippi and eastern Louisiana. Al- though it is not yet known whether the species range extends beyond the eastern USA, given the known ranges of Atlantic coastal plain plant species (Sorrie and Weak- ley, 2001) the sand plains of southern Nova Scotia would appear to be an appropriate location for future surveys. Preferred Habitats: Vertigo malleata occurred in ap- proximately two-thirds of all surveyed acid habitats. In southern Maine and Massachusetts it was found in damp and lightly compacted leaf litter on Sphagnum bogs un- der a dense cover of ericaceous and other acidophile shrubs (e.g., Gaylussacia, Vaccinium, Kalmia, and Myrica). In this region it was also present in Atlantic white cedar bogs, where it occurred in leaf litter accu- mulations on mossy hummocks, In the New Jersey Pine Barrens V. malleata was found in dense leaf litter under tall heath (Vaccinium, ach pas ia, Kalmia), Myrica, and Ilex scrub at the edges of bogs, Atlantic white cedai swamp forest, and mesic microsites in upland pine-oak forest. Populations in North and South Carolina were primarily located under dense heath, bay, holly, and wax myrtle scrub in pocosins, bay forest, wet-mesic pine woodland, and pine-wiregrass savanna. At Pocosin Lakes National Wildlife Re fuse, for example, V. malleata was abundant in leaf litter on scrub se cee within flooded pond pine woodland (Table 1, site 55), absent in adjacent broadleaf woodland, and is sent only in rela- tively low numbers at the transition zone (site 56). Popu THE NAUTILUS, Vol. 121, No. 1 Figures 1-19. Vertigo malleata and related taxa. 1-4. Vertigo malleata. Holotype, NMW.Z.2005.011.03530, Holly Shelter Game Lands, Pender County, North Carolina, 34°31'57" N, 77°44'41" W; 1. Apertural view. 2. Abapertural view. 3. View showing parietal and upper palatal lamellae. 4. Sculpture on body whorl surface, width of detail is 0.25 mm. 5. Vertigo malleata, second specimen from the type locality, NMW.Z.2005.011.03831, showing more conical shell shape. 6. Vertigo malleata, NMW.Z.2005.011.03832, Stafford Forge WMA, Ocean County, New Jersey, 39°42'44" N., 74°22'10" W, showing small size and lack of infraparietal and subcolumellar- basal lamellae. 7-8. Vertigo malleata, NMW.Z.2005.011.03533, Wells Heath, York County, Maine, 43°20'2"” N, 70°38'24" W. 7. \pertural view. 8. View from apex showing apical whorls and the basal apertural dilation. 9. Vertigo malleata, NMW.Z,2005.011.03834, Wilma Station, Liberty County, Florida, 30°9'34" N, 84°57'39" W, showing strong shell sculpture, a series of subcolumellar-basal lamellae, and a weak infraparietal lamella. 10, 11. Vertigo malleata, NMW.Z.2005.011.03835, Pond Creek seep, Covington County, Alabama, 31°6'12"” N, 86°32'3" W. 10. Apertural view showing subcolumellar and basal lamellae, an indistinct nodule below the columellar lamella, and lack of an infraparietal lamella. 11. Abapertural view, 12. Vertigo malleata, NMW.Z.2005.011.03536, Collins Bay, Ware County, Georgia, 31°05'12" N, $2°36'56" W, showing elongate shape, fused sub- columellar and basal lamellae, and distinct sinulus. 13. Vertigo malleata, NMW.Z.2005,011.03837, Skunknett Audubon Preserve 2, Barnstable County, Massachusetts, 41°38'59" N, 70°22'31" W, showing light shell color and basal lamella only. 14. Vertigo malleata, NMW.Z.2005.011.03838, Peterson Swamp WMA, Plymouth County, Massachusetts, 42°00'37" N, 70°49'4"” W, showing small size, vestigal infraparietal, and reduced basal lamellae. 15. Vertigo malleata, NMW.Z.2005.011.03839, Lewis Ocean Bay Preserve, Horry County, South Carolina, 33°47'16" N, 75°50'56" W, showing bi-lobed basal lamella. 16. Bothriopupa tenuidens (C.B. Adams, 1845), FMNE 106420, Louis Brand Collection, Columbia University. 17. Bothriopupa conoidea (Pfeiffer, 1853), FMNH 119055, Kyk- Over-All Island, Kartabo, British Guiana. 18. Vertigo conecuhensis, JON 12364, Pond Creek seep, Covington County, Alabama, 31°6'12" N, 86°32'3" W. 19. Vertigo alabamensis, JCN 10781, Lanier Quarry, Pender County, North Carolina, 34°37'49" N, 77°40/27" W lations in Georgia, Florida, and Alabama were found pri- marily in bay scrub along swamp margs small water courses, and seepage Zones within pinelands Populations were also rarely encountered in mesic pine forest frag- ments that had escaped frequent fire management (see below): again, individuals were restricted to humid litter tumulations. Throughout its range, Vertigo malleata ppeared to avoid even moderately less acidic habitats uch a sedge meadows (Maine, Massachusetts), cattail swamps and marshes (Maine, Massachusetts, New Jer- sey), and bottomland bald cypress/water tupelo/ sweetguin forests (North and South Carolina, Georgia, Florida, and Alabama) Associated Land Snails and Community Composi- tion: Across all 49 analyzed sites (Table 2), Vertigo malleata constituted 35% of total individuals. This frac- tion appeared to be inversely correlated with latitude, B. F. Coles and J. C. Nekola, 2007 g2°W ee 88°W 84°W 96°W g2°W 88°W igure 2 Figure 20. 80°W 76°W 84°W Distribution of Vertigo malleata in eastern North America. Black circles represent sites supporting populations and 72°W 68°W 42°N 38°N 34°N 26°N 80°W 76°W 72°W open circles represent inventoried sites that do not harbor this species. ranging from 17% in the New Jersey Pine Barrens to 32% in New England, 35% in the Carolina coastal plains, and 76% in the Gulf coastal plain. Population de ioe s of Vv _malle vata were frequently observed to exceed 500 per m*, with an estimated de ue of the order of 2000 per m- at the type locality (Table 1, site 51). These densities range among the highest re ets ed for any land snail spe- cies (Frest and Johanne s 1995, Cameron 2003). A total of 34 terrestrial mollusk taxa and 5886 indi- viduals were observed from these sites (Table 2). The average number of co-occurring taxa was 3.69 + 0.37, and ranged from 0-9. Throughout its range, the ten most frequently co-occurring taxa were: Striatura milium 17% of all other individuals), Strobilops texasiana (15%), Vertigo milium (13%), Punctum minutissimnaun (11%), Vertigo alabamensis (9%), Striatura meridionalis (8%), Gastrocopta pentodon (8%), Euconulus trochulus (4%), Euconulus chersinus (2%), and Gastrocopta con- tracta (2%). The most frequent co-occurring taxa varied by region: Striatura milium and Punctum minutissiu- mum in New England: Striatura meridionalis, Striatura milium, Punctum minutissimum, and Gastrocopta pent- odon in the New Jersey Pine Barrens; Strobilops texasi- ana, Vertigo milium, and Vertigo alabamensis along the Carolina coastal plain; and Vertigo alabamensis, Stria- tura meridionalis, and Gastrocopta pentodon along the Gulf coastal plain. Co-occurring Vertigo taxa also varied by region: Vertigo cristata, Vertigo perryi, and Vertigo Page 26 THE NAUTILUS, Vol. 121, No. 1 Table 2. Frequency of co-occurring species across the range of Vertigo malle ata! { Number of co-occurring individuals (%) New Jersey Carolina coastal plain Gulf coastal plain Total Taxon New England Vertigo malleata sp. nov. 308 Striatura milium (Morse, 1859) 480 Strobilops texasiana Pilsbry & Ferris, 1906 Vertigo milium (Gould, 1840) Punctum minutissumum (1. Lea, 1541) 113 Vertigo alabamensis Clapp, 1915 Striatura meridionalis (Pilsbry & Ferris, 1906) Gastrocopta pentodon (Say, 1821) 6 Euconulus trochulus (Reinhardt, 1883) Euconulus chersinus (Say, 1821) Gastrocopta contracta (Say, 1822) Glyphyalinia solida (H. B. Baker, 1930) Glyphyalinia sp. Vertigo oralis Sterki, 1898 Hawaiia miniscula (A. Binney, 1840) Glyphyalinia luticola Hubricht, 1966 Vertigo conecuhensis Clapp, 1915 Gastrocopta tappaniana (C. B. Adams, 1842) 4 Zonitoides arboreus (Say, 1816) 12 Euconulus fulvus (Miiller, 1774) 20 Vertigo ovata Say, 1822 Vertigo ovata Say, 1822 Neohelix solemi Emberton, 1988 ] Ventridens cerinoideus (Anthony, 1865) Vertigo cristata (Sterki, 1919) 13 Nesovitrea electrina (Gould, 1541) 6 Vertigo perryi Sterki, 1905 5 Gastrocopta riparia Hubricht, 1978 Helicodiscus parallelus (Say, 1817) Troidopsis soelneri (J. B. Henderson, 1907) Deroceras sp. Glyphyalinia indentata (Say, 182° | Vertigo rugosula Sterki, 1890 Vertigo ventricosa (Morse, 1865) 2 Striatura ferrea Morse, 1864 1 Triodopsis hopetonensis (Shuttleworth, 1852) Total co-occurring individuals 664 Co-occurring species richness 13 340, 1S07 779 3234 407 119 1006 61 $35 ll 907 765 768 398 164 675 494 64 558 413 25 46 484 258 156 45 465 216 216 120 3 123 109 109 15 64 19 98 28 63 91 61 6 67 53 53 28 12 3 43 39 39 28 32 5 14 31 20 20 20) 20 1S 19 19 19 13 6 5 3 3 3 3 3 3 2 2 | 2 2 2 2 1 | 1 1613 3359 250, 5886 9 26 LO 34 ' Data taken from 49 discrete sites of the junior author collection (lots prefixed by JCN in Table 1), * Juveniles and young adults of unclear identity. ventricosa were sympatric in New England, while Ver- tigo milium, Vertigo alabamensis, Ve rtigo oralis, Vertigo conecuhensis, and Vertigo rugosula were sympatric in the Carolina and Gulf coastal plains. CONSERVATION IMPLICATIONS The data presented here show that Vertigo malleata is a characteristic component of the base-poor biota of the Atlantic and Gulf coastal plains, having been found in 66% of surveyed acid sites, and accounting for up to 75% of all mollusks in these sites. The sundance ancl wide- spread occurrence of V. malleata would seemingly sug- gest that it is not of immediate conservation concern. However, it appears that the species is in fact under threat because of the widespread use of fire as a man- agement tool. Many coastal plain habitats, including those of V. malleata (i.e., pine woods, pine-wiregrass sa- vanna, and pine barrens), have come to be wiewed by many plant ecologists as pyrogenic (Myers, 1985; Chris- tensen, L98S) \ and are being typically managed by anthro- pogenic fire return inte eal of <5 years, with mz uny areas being burned annually. However, such high- freque ney fire man: wement policies have been tas to exact a pan negative impact on total biodiversity, including Le pidopter: i, Homoptera, Hymenoptera, Araneae, ( ‘ol- lembola (Swengel, 1996, 1998: Harper et al., 2000), and terrestrial Mollusca (Ne kola, 2002b). The impact of fire on Vertigo malleata is illustrated by its distribution in the Appi lachicola uplands of F lorida. B. F. Coles and J. C. Nekola, 2007 Page 27 We were unable to find V. malleata in forest that had been burned within three years, however, the presence of substantial populations in a tiny unburned inholding of mesic pine forest (Table 1 site 11), unburned mesic pine- red maple-Atlantic white cedar forest (Table 1 site a and unburned mesic margins of wetlands (Sites 8, 12-13, 15-17) suggest that while it is not ay a restricted to wetlands, it has become lar gely limited to these sites simply because they remain unburned. While these observations require further inv estigation, we esti- mate that at least 95% of the \ the Apalachicola National Forest has been eliminated by management practices. Conversely, the presence of 14 malleata in mesic bay-pine forest that had regenerated after burn (Table 1 site 10) shows that, apart from its intrinsic interest as an extreme calcifuge, this snail can potentially be used to monitor recovery from over- burning. ACKNOWLEDGMENTS We thank Beata Pokrysko (Museum of Natural History, Wroclaw University, Wroclaw, Poland) for comments concerning the identity of Vertigo malleata as an unde- scribed species, and to Jochen Gerber for loans of Both- riopupa material from the FMNH collection. José H. Leal, Barry Roth, and John Slapcinsky all provided useful comments on earlier drafts. The following U. S. Federal Government agencies, U.S. State Gov ernment agencies, private organizations and individuals have helped during this study by providing access to lands and information on sites: Alabama Department of Conservation and Natural Resources, Forever Wild Program (Eric Soe- hren); Maine Department of Inland Fisheries and Wild- life. Wildlife Resource Assessment Section (Beth Swartz): Massachusetts Natural Heritage and Endan- gered Species Program (Tom French and Jennifer Loose); New Jersey Department of Environmental Pro- tection, Division of Fish and Wildlife, Endangered Spe- cies Program (Dave Golden); New Jersey Department of Environmental Protection, Division of Parks and Forests (Thomas Keck, and the Superintendents of Belleplain State Forest-W. Scott Mauger, Brendan T. Byrne State Forest-Christian Bethmann, and Wharton State Forest— Lynn Fleming); North Carolina Division of Forestry Re- sources, Bladen Lakes State Forest (Michael Chesnutt); North Carolina Wildlife Resources Commission (Stephen Hall and Randall Wilson); South Carolina De- partment of Natural Resources, (Lewis Ocean Bay Pre- serve Manager—Jamie Dozier); The Nature Conservancy, North Carolina Chapter (Dan Bell and Bruce Sorrie): The Nature Conservancy, Southern Maine Field Office (Parker Schuerman and Keith Fletcher); U. S. Forest Service. Apalachicola National Forest, Wakulla District (Marcus Beard): U.S. Fish and Wildlife Service, Pocosin Lakes National Wildlife Refuge (Wendy Stanton). V. malleata population of LITERATURE CITED Baker, F. C. 1939. Fieldbook of Illinois land snails. Illinois Natural History Survey, Manual 2. Illinois Natural History Survey Division, Urbana, 166 pp. Boycott, A. E. 1934. The habitats of land mollusea in Britain. Journal of Ecology. 22: 1-38. Burch, J. 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Academy of Natural Sciences, Philadelphia, pp. 521-1113 + xlvii. ? ere Page 28 THE NAUTILUS, Vol. 121, No. 1 Pokryszko, B. M. and R.A.D. Cameron. 2005. Geographical variation in the composition and richness of forest snail faunas in northern Europe. Records of the Western Aus- tralian Museum, Supplement 68: 115-132 Schilthuizen, M., H. N. Chai, T. E. Kimsin, and J. J. Ver- meulen. 2003. Abundance and diversity of land snails (Mollusca: Gastropoda) on limestone hills in Borneo. Raffles Bulletin of Zoology 51: 35-42. Sorrie, B. A. and A. S. Weakley. 2001. Coastal plain vascular plant endemics and phytogeographic patterns. Castanea 66: 50-82. Swengel, A. B. 1996. Effects of fire and hay management on the abundance of prairie butterflies. Biological Conserva- tion 76; 73-85. Swengel, A. B. 1998. Comparisons of butterfly richness and abundance measures in prairie and barrens. Biodiv ersity and Conservation 7: 639-659. ° Turgeon, D. D., J. F. Quinn Jr., A. E. Bogan, E. V. Coan, F. G. Hochberg, W. G. Lyons, P. M. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F. G. Thompson, M. Vecchione, and J. D. Williams. 1998. Com- mon and Scientific Names of Aquatic Invertebrates from the United States and Canada, Mollusks, 2nd edition. American Fisheries Society, Special Publication 26. American Fisheries Society, Bethesda, ix + 526 pp: THE NAUTILUS 121(1):29-36, 2007 >a0e@ 29 Page 29 Population dynamics of the fingernail clam Sphaerium occidentale (Lewis, 1856) Genes Sphaeriidae) in an intermittent pond Meghna Roy D. Dudley Williams Department of Life Sciences University of Toronto at Scarborough 1265 Milit: uv Trail, Scarborough Ontario MIC 1A4, CANADA ABSTRACT A population of Sphaerium occidentale (Lewis, 1856) was stud- ied over a two-year period in an intermittent freshwater pond in southern Ontario, Canada. Sub- populations in control areas of the pond showed marked differences between the two years, which appeared to be related to different hydroperiods (34 days in 2001 vs. 94 in 2002), water temperature, and density of suspended bacteria. In the first year, there was recruitment of young into the population in April/May, but there appeared to be very little or no Reproduction in the second year. Resource (decaying riparian leaves) addition and remoy al experiments performed in the second year had an impact on some but not all of the physico-chemical parameters measured in the pond, and also affected bacterial densities. However, there were no significant overall effects of these manipulations on sub- populations of S. occidentale, although there was a trend to- ward greater density and biomass in the resource removals during the first half of the hydroperiod. Sphaerium occidentale appears to be well adapted for survival in such harsh environ- ments via a specialized physiology ry and, reproductively, employ- ing a bet-hedging strategy However, populations may occa- sionally be reduced to le vale that may result in local extinction. To counter the latter, sphaeriids have dispersal mechanisms that allow recolonization from metapopulations. INTRODUCTION Sphaeriid clams are widely distributed and many specie S inhabit temporary fres shwater habitats. They are impor- tant components of the benthic communities of such habitats where they act as water clarifiers and organic nutrient sinks (Thorp and Covich, 2001). Sphaeriids are known to be simultaneous hermaphrodites, with the abil- ity to self-fertilize. a trait well suited to founding and maintaining populations in temporary waters (W sThanae: 2006). Sphaerium occidentale (Lewis, 1856), commonly Corresponding author: Meghna Roy email: meghna@zoo.utoronto.ca known as Herrington’s fingernail clam, is unique among the Sphae sriidae in that it is exclusive to temporary waters (McKee and Mackie, 1951). Along with all other sphaeri- ids, individuals of this species brood its direct-developing young within special sacs in the immer demibranchs of the gills, from where they are eventually released as benthic juveniles (Mackie et al., 1974). Although many sphaeriids are synchronous brooders, species of ‘Sphae rium are se- quential brooders—that is, several sets of embryos, each ina separate stage of ontogeny, are present within the brood sacs, together, each the product of a separate spawning (Mackie, 1978). Presumably, this trait is a bet- hedging strategy conducive to releasing a subset of viable young rapidly at the beginning of a hy droperiod of un- certain length, while maintaining a reserve should the first subset ie lost (Stearns 1992). Although the popula- tion dynamics of Sphaerium occidentale are not well known, McKee pe Mackie (1981) observed that the species completed its life cycle in 24 weeks when main- tained under a stable hydroperiod in the laboratory, as opposed to three years in an intermittent pond. This finding supports the hypothesis of Thorp and Covich (2001) that seasonal variation in water level drives bivalve life history traits. The purpose of the present study was to determine any relationship between hy droperiod length and the popul ition dynamics of S. occidentale in an in- termittent pond, and to explore other possible influences through field manipulation experiments. MATERIALS AND METHODS The study population lives in an intermittent pond in Vandorf, southern Ontario, Canada. The pond is devoid of fish and has an area of approximately 1000 mn at the beginning of hydroperiod. It is surrounded by a hay field ane wieed deciduous woodland and supports emergent vegetation comprising mainly Phalaris grass. The pond substrate is primarily muddy and homoge neous through- out. The population was studied over a two year pe riod Page 30 THE NAUTILUS, Vol. 121, No. 1 in 2001 the pond held water for 34 days and had a maxi- mum depth of 66 cm; in 2002 it held water for 94 days and had a maximum depth of 48 cm. The populations were studied in six randomly chosen areas of the pond that were enclosed with watertight, circular galvanized sheet-metal walls installed prior to snowmelt in 2001. The walls were embedded 10 cm into the substratum and rose well above the water surface, forming corralled homogeneous sections of the pond each 2.4 m in diameter. Two of the enclosures were chosen at random to serve as controls. In the autumn of 2001, the locations of the control enclosures within the pond were changed in order to avoid carry-over effects, and the other four enclosures were manipulated to either receive additional riparian leaf litter (two each received a mixture of dry maple and beech leaves |~3.0 kg] that had been dried for 24 hours at 100°C), or have litter removed (~1.540.04 kg of leaves and other vegetation from each of the remaining two). It might be argued that the use of imperforate galvanized metal enclosures may have ren- dered the treatments uninhabitable (i.e., produced a container effect) for S. occidentale, but that is highly unlikely as clams are virtually immobile and non- selective filter feeders. Further, evidence from other taxa (e.g., ciliates) in this pond showed that the enclosures resulted in higher species richness and abundance (An- drushchyshyn et al., 2006). The environments within the enclosures were sampled on a weekly basis beginning as soon as the pond basin filled in 2001 and again in 2002. The following param- eters were measured: water depth, with a one-meter stick, and dissolved oxygen, pH, temperature, and con- ductivity with a portable Hach Kit spectrophotometer (DR2000: Hach Company, Loveland, Colorado, USA). Turbidity, ammonia, nitrate, and phosphorus were mea- sured using a Hydrolab multiprobe (Hydrolab Corpora- tion, Austin Texas, USA), and chlorophyll ad was mea- sured using the acetone extraction method (American Public Health Association, 1995). To assess bacterial density, a column of water extending from the bed to the surface was collected using a plastic tube; this was done at two locations within each enclosure and then com- bined in a bucket from which a subsample of 30 cm Was removed and stained with acridine orange. These stained samples were then filtered through a 0.2 wm Sartorius filter (Cat. No. 13007) and the bacteria counted (on ~20 fields with a concentrations of ~200 bacteria per field of view) under an epifluorescence microscope, following the method of Sorokin (1999). ( Q uns Were Se imple d using a calvanized-steel box sam- pler (area 0.1 m~) which was inserted 2 cm into the pond Bed. Two such bee samples were taken at regular intervals from random locations in the control enclosures in 2001 and from the control and treatment enclosures in 2002. All samples were put in large Ziploc bags, labeled, and preserved in 4% formalin in the field. In the labo- ratory, clams were removed by sieving and handpicking inder a magnifying lens from samples collected on 7 April, 3 May, 10 July, and 21 August in 2001, and 7 April, 6 May, 3 June, 11 July, and 6 September in 2002. All specimens were counted, blotted on filter paper to re- move excess water, and weighed to the nearest 0.0001 ¢ using a digital balance. All specimens proved to be Sphaerium occidentale, and identification was confirmed by Dr. Gerald Mackie (University of Guelph, Canada). Clam le ngths (anterior to posterior) were measured to the nearest 0.1 mm using a dissecting microscope fitted with an ocular micrometer, and specimens were assigned to the following four length classes: <2.5, 2.5-4.0, 4.1— 5.5, and > 5.5 mm. RESULTS CONTROL POPULATIONS: In 2001, the hy ‘drope riod of the Vandorf pond was short (34 days, from 7 April to 17 May) and likely related to a rapid 10° - warming of the pond i in the first week of May (Figure la, b) and low rainfall. Associated with this warming was a significant increase in the density of sus- pended Thactenia [Figure le; F=7.14, p=0.02 . from a re- peated-measures ANOVA], perhaps partly a concentra- tion effect, and a marked increase in the growth of indi- vidual clams (Figure 2a). From May to July, 2001, mean biomass increased from 0.33+0.02 to 2.1+0.22 ¢ 0.1 m 2 of pond bed area (ANOVAR F=5.95, p=0.03). At the times of sampling the dry pond bed in July and Septem- ber, 2001, no additional growth was noted. In 2002, the hydroperiod was considerably longer (94 days) and started almost three weeks earlier (18 March to 19 June; Figure 1b), While water temperature attained the same maximum value as in 2001, its increase was more gradual (Figure la). Bacterial densities throughout 2002 were more stable, except for a minor increase in early June (Figure le). Clam growth during 2002 was very different from 2001 and was largely limited to shifts from size class | to 2 in May-June (Figure 2b; ANOVAR F=12.88, p=9.005). Biomass increase -d from 0.24+0.14 ¢ 0.1m in May to 0.88+0.39 ¢ 0.1 m~ in July (ANOV AR F=15.13, p=0.0001). a Between-year comparisons suggest that while there was recruitment of young into the population in April/ May of 2001 (Figure 2a), the similarity in population structure between Se ptember, 2001 and April/May, 2002 and concomitant decline in density of juveniles (Figure 2b) indicate that re production did not occur im 2002. MANIPULATED POPULATIONS: The resource addition and removal experiments per- formed in 2002 had an impact on some but not all of the physico-chemical parameters measured in the Vandorf pond. Largely unaffected, compared with the control en- closures, were water temperature (although there was an increase in the resource addition enclosures in May), water depth, pH phosphorus, and chlo- rophylla. Affected were dissolved oxygen levels (typically (Figure 3 a,b,c) M. Roy and D. D. Williams, 2007 Page 3] oO —@— 2001 oo 15.4 ——m— 2002 = 2 g Sos], 0) T 1 1 T [—— r T 1 March April May June Month b) Depth (cm) March April May June Month Cc) s 30000 - w 25000 - i) xe} E 20000 - © 15000 - & 5 10000 - S 4 S 5000 2 (0) TT oer ee a oe Mar-18 Apr-1st Apr-3rd May-1st May-2nd Jun-03 Jun-20 week week week week Date Figure 1. Seasonal variation in water temperature, depth, and bacterial density in the control enclosures in 2001 and 2002. Mean values are shown for the control enclosures. Error bars for bacteria indicate +1SE (n = 2). Page 32 THE NAUTILUS, Vol. 121, No. 1 Control - 2001 No we sample taken (7) (40) (448) (363) |r| ol} Control - 2002 a ive = 4 (69) (39) (83) (87) (96) Size class a= rej asx — (79) (54) (99) (167) (123) Resource removal - 2002 tet dt (128) (114) (188) (145) ori May janet (aii guy ay) September ( (dry) s/n) ol) Figure 2. Seasonal length frequency a aneee of Ps Sphaerium occidentale in 2001 and 2002. Size classes 1, 2, 3, and 4 represent clams < 2.5, 4, 5.5, and > in length. Numbers in parentheses are sample Sizes. 5.5 mm, respe ative ly, highest in the resource removal and lowest in the re- source addition treatments) and nitrate (highest in the resource additions early on in the hydrope wig) 4 ab). Bacterial densitie Ss were highest in the resource Seen treatments, lowest in the resource removals, and intermediate in the controls during the first half of the hydroperiod, but more similar thereafter (Figure 4c) Despite differences in the above parameters, there Figure S were no significant overall treatment effects on the den- sities or biomass of S. occidentale, although there was a trend towards greater density and biomass in the re- source removals during the first half of the hydroperiod (Figure 5a,b). Neither did there appear to be any notable differences in the population structures in the addition and removal enclosures compared with the 2002 controls (Figure 2b,c,d). DISCUSSION a sharp rise in water temperature and rapid decline in le pt th of the Vandorf pond in early May of 2001, accom- panie od by low rainfall, likely promote 1d the sion hydro- pe riod in that year. Bivi alve feeding filtration rates are known to be proportional to the ambient temperature | horp and ¢ Lame 8 2001) ing of the pond fortuitously allowed Thus the sudden 10°C warm- occidentale to benefit maximally from the high density of suspended bacteria available at that time, and to grow rapidly. How- er, and in spite of the longer (~3 times) hydroperiod in 2002, during this second year S. occidentale individuals grew much more slowly—although a growth spurt did occur during the higher temperatures "reached prior to pond dry-up in June. Based on this two-year comparison, it would appear that individual clam growth does not necessarily benefit from a longer hy rope »riod—indeed a shorter one may be better provided that food supply and water temperature are favorable. Thomas (1963) noted, however, that S. partumeium (Say, 1822) grew more un- der a longer hydroperiod although its life span is 12-13 months compared with up to 3 years in S. occidentale. The former species may well be predisposed to contin- ued growth in that its semelp: wrous strategy commits it to development and reproduction under sometimes less than optimal pond conditions. S. occidentale appears to demonstrate more plasticity in its growth rate (McKee and Mackie, 1981), and its iteroparous strategy may allow it, on occasion, to forego a less than favourable repro- ductive opportunity, Only adult (our size class 4) S. occidentale release off- spring (McKee and Mackie, 1951). Some of the sub- adults present in April 2001 grew rapidly, attained size class 4, and produced large numbers of size class 1 and 2 young, as evident from the very high densities found in July 2001. As S. occidentale is, as previously noted, a sequential brooder it is conceivable that the high July densities may have resulted from multiple release events of young from ¢ April until the end of the hydroperiod. The same size-class proportions present in the July and September samples indicate that no further growth took place during the period when the pond bed was dry, which agrees with the findings of Mckee and Mackie (1983) for this species. Further, the presence of the same distribution of size classes in April (2002) shows that there was no winter growth or container effect from the use of imperforate metal enclosures. The substantial drop in clam density between September 2001 and April 2002 likely indicates winter mort: lity. Throughout 2002 no significant increase occurred in the control population and this, together with modest shifts towards the large size classes by September, suggests that the population largely | failed to breed in this second year. Unlike S. partumeium, which is more commonly found in permanent waters (Clarke, 1973), and has a semelpa- rous, univoltine life cycle where only juveniles aestivate and the resulting adults breed and die the next spring (Way et al., 1980), S. occidentale can aestivate in all size classes and has the potential to breed several times and over more than one hydroperiod as pond conditions al- low. Interestingly, the initial samples collected in a 2001 produced very few clams, and no live adults. possible explanation for this is that when sphaeriids en their terminal size, they do not survive a subsequent dry phase (Way et al. 1980). It is like ly, then that the majority of the previous year’s population had been adults at the end of the 2000 hydroperiod, The popula- tion was thus at a very vulnerable stage in early 2001, but M. Roy and D. D. Williams, 2007 Page 33 a) —@— Resource addition O ——l—— Resource removal sill Control Qa 5 -E — oO ww s Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun- 18 25 31 7 16 22 29 6 13.20 27 03 10 17 Date Depth (cm) Mar- Mar-Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun- Jun- 18 25 31 7 16 22 29 6 13 20 27 £403 10 17, = 20 Date 8 7 6 = 2) a4 3 2 1 0 Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun- 18 25 31 7 16 22 29 6 13 20 27 03 10 17 Date Figure 3. Seasonal variation in water temperature, depth, and pH in the treatment enclosures in 2002. Mean values are shown for the control. resource addition, and resource removal enclosures. Page 34 THE NAUTILUS, Vol. 121, No. 1 Dissolved 0 (mg/L) Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun- 18 25 31 ig 16 22 29 6 13 20 27 #203 #10 = «17 Date b) Nitrate (mg/L) Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun- 18 25 31 7 16 22 29 6 13 20 27 #03 10 17 Date a) —@— Resource addition = ——™—— _ Resource removal é Control 2 Ee =) e & © oO 19) fan) Mar-18 Apr-7 Apr-22 May-06 May-20 Jun-03 Jun-17 Date Figure 4. Seasonal variation in oxygen, nitrate, and bacterial densities in the treatment enclosures in 2002. Mean values are shown for the control, resource addition, and resource removal enclosures. Error bars for bacteria indicate +1SE (n = 2). was quite quickly restored by July, only to suffer sub- ated with an increase in dissolved oxygen and a lower stantial winter mortality in all size classes later that same concentration of suspended solids, although improved yeal water clarity did not affect chlorophyll @ production. Re- Removing resources from the enclosures was associ- source addition reduced oxygen levels and increased tur- M. Roy and D. D. Williams, 2007 a) x Control = = Resource addition oO — Resource removal > — 7p) c o a) = ase, O April May a = Se jo) ™~ ie) ~~ Yn n © = 2 jaa) = Ba O April May June July (dry) September (dry) Month July (dry) September (dry) Month Figure 5. Mean values of Sphaerium occidentale densities in numbers 0.1 m~ and biomass in g 0.1 m~ in the treatment enclosures in 2002. Mean values are shown for the control, resource addition, and resource removal enclosures. Density and biomass bars indicate +1SE (n=2) and +1SD (n=2), respectively. bidity, likely as a result of suspension of more litter breakdown products. While it might be expected that an increase in suspended food particles (including bacteria in the first half of the hydroperiod) would have benefit- ted the S. occidentale population (as shown for other detritivores, by Richardson, 1991), it is known that too high a concentration of suspended materials can inter- fere with the filtering and respiratory mechanisms of freshwater bivalves—although sphaeriids are known to be very tolerant of hypoxia (Thorp and Covich, 2001). Further, sphaeriids can also feed on pond bed deposits and thus a direct response to food particle manipulation may not be detectable. Despite the observed differences in the physico-chemical environments and food levels between the treatments and the controls, there were no apparent differences in the structure (size and growth) of the sub-populations. Lack of a differential response may be a reflection of the fact that intermittent ponds are normally subject to large within-vear and between-year fluctuations in environmental variables (due to dilution, . and thus their inhabitants may not show population response patterns typical of those evaporation, etc known from permanent ponds (Brénmark and Hansson, 1998S). The responses of the natural and manipulated sub- populations of S. occidentale in the Vandorf pond suggest that length of the hydroperiod was the major driving force on population dynamics. Further, the species ex- hibits a number of important adaptations that enable population survival despite large inter-year variations in its habitat, although populations may occasionally be reduced to dangerously low numbers that may result in local extinction. Sphaeriid clams are known, however, to be able to colonize/recolonize these small ponds via transportation on the bodies of more-mobile pond inhabitants (e.g., Fryer 1974). It is likely that such re- colonization events from a regional metapopulation are common. ACKNOWLEDGMENTS We thank the Natural Sciences and Engineering Re- search Council of Canada for funding, and Katarina Magnusson for collecting the samples Page 36 THE NAUTILUS, Vol. 121, No. 1 LITERATURE CITED American Public Health Association. 1995. Chlorophyll. In: Franson, M. A. H. (ed.) Standard Methods for the Exami- nation of Water and Wastewater. American Public Health Association, American Water Works Association and Wa- ter Environment Federation, Washington, pp. 10.17— 10,24. Andrushchyshyn, O. P., A. kK. Magnusson, and D. D. Williams. 2006. Responses of intermittent pond ciliate populations and communities to in situ bottom-up and top-down ma- nipulations. Aquatic Microbial Ecology 42: 293-310. Brénmark, C. and L. Hansson. 1998. The ‘biology of ihe and ponds. Oxford University Press, New York, 216 pp. Clarke, A. H. 1973. The freshwater molluses of the Canadian Interior Basin. Malacologia 13: 1-509. Fryer, G. 1974. Attachment of bivalve molluscs to corixid bugs. Naturalist 28: 18. Mackie, G. L. 1978. Growth dynamics in natural populations of Sphaeriidae clams (Sphaerium, Musculium, Pisidiwm). Ca- nadian Journal of Zoology 57: 441-456. Mackie, G. L., S. U. Qadri, and A.H. Clarke. 1974. Develop- ment of brood sacs in Musculium securis Bivalvia: Sphaeri- idae. The Nautilus 88: 109-111. Mckee P. M. and G. L. Mackie. 1981. Life history adaptations of the fingernail clams Sphaerium occidentale anil Muscu- lium securis to ephemeral habitats. Canadian Journal of Zoology 59: 2219-2299, McKee P. BE and G. L. Mackie. 1983. Respiratory adaptations of the fingernail clams Sphaerium occidentale and Muscu- lium securis to ephemeral habitats. Canadian Journal of Fisheries and Aquatic Sciences 40: 783-791. Richardson J. S. 1991. Seasonal food limitation of detrivores in a montane stream: an experimental test. Ecology 72: 873— SS7. Sorokin, Y. I. 1999. Aquatic microbial ecology—a textbook for students in environmental science. 99. Backhuys Publish- ers, Leiden, 248 pp. Stearns, S.C. 1992. The Evolution of Life Histories. Oxford University Press, Oxford, 249 pp- Thomas, G. J. 1963. Study of a population of sphaeriid clams in a temporary pond. The Nautilus 77: 37-43. Thorp J. H. and A. P. Covich. 2001. Ecology and Classification of North American Freshwater Invertebrates. 2°¢ Edition. gry Press, New York, 1056 pp. Way, C. M., D. J. Hormbach, and A. J. Burky, 1980. Com- pe a life history tactics of the sphé veriid clam Muscu- lium partumeium (Say), from a permanent pond and a temporary pond. American Midland Naturalist 104; 319— 397 Williams, D. D. 2006. The Biology of Temporary Waters. Ox- ford University Press, Oxford, 337 pp. a SSS SSS SSS SSS SSS SSS THE NAUTILUS 121(1):37-39, 2007 Page 37 Research Note Paleoenvironmental significance of the eastern mud snail, Ilyanassa obsoleta (Say, 1822 ), from a microtidal coastal sequence of southern New Eneland Ilya V. Buynevich Woods Hole Oceanographic Institution Geology and Geophysics Department, MS#22 Woods Hole, MA 02543 USA ibuynevich@whoi.edu Reconstruction of former sea-level positions is one of the key issues in Quaternary paleoenvironmental research. Along with high-marsh peat that has been widely used for regional sea-level reconstructions due to its narrow and robust elevation range, intertidal mollusks inhabiting protected coastal embayments must also be eonsideied as compleme ntary reference points for the water levels. This paper presents evidence for potential use of the eastern (Atlantic) mudsnail Iyanassa obsoleta (Say, 1822), as a paleoenvironmental indicator in middle Ho- locene deposits of southern New England. The eastem mudsnail occupies tidal flats along the Atlantic seaboard, often occurring in dense populations on the sediment surface of intertidal mudflats or mixed flats (Brenchley, 1980; Whitlatch, 1952; Culbert and Ra- leigh, 2001). Their feeding habits include scavenging, predation, algal foraging, por deposit-feeding (Weiss, 1995: Kelaher et al., 2003), with feeding svthans shown to be correlated with tidal cycles ( Goberson: 1979). Na- tive along the Atlantic seaboard, Ilyanassa obsoleta has been recently under stress of competitive exclusion and egg predé ition by invasive gastropods, such as Littorina littorea (Brenchley, 1982: Carlton, 1992) and has itself become an invasive species in some parts of the West Coast (Race, 1982). Although these snails are known to burrow into the substrate or migrate into shallow subtidal depths during the winter (Batchelder, 1915; Dexter, 1961; Brenchley, 1980), the depth of migration is likely to be limited bv tidal range and wave energy. For ex- ample, in a microtidal coastal setting (mean tidal range: 0-2 m) where tidal flats are fronted by wave-dominated coastal barriers, the vertical habitat range of I. obsoleta will be relatively narrow (Whitlatch, 1982). This fact, combined with occurrence of the eastern mud snail in post-glacial deposits (past 10,000-15,000 years; Carlton, 1992), makes this species a potential indicator of former sea level. The vertical range of a particular plant or animal spe- Corresponding author: Ilya Buynevich email: ibuynevich@whoi.edu cies preserved in a geological record and its position relative to a specific tide len el (indicative meaning; van der Plassche, 1986; Donnelly et al., 2004) can he esti- mated based on modem ecological response of that or- ganism to tidal inundation. Therefore, comparison of in- situ shells of I. obsoleta with an adjacent reliable sea-level indicator, such as high-marsh peat, is the first step in assessing its paleoenvironmental significance. The aim of this paper is to use similar ages of L. obsoleta and high- marsh peat in submerged banlianrer deposits offshore Martha's Vineyard wane, Massachusetts, as evidence of a potential use of this gastropod as an independent sea- level indicator. The microtidal barrier coastlines of southern New En- gland and Long Island have been formed in a regime of post-glacial marine transgression, with Holocene sedi- mentary sequences now submerged on the inner conti- nental shelf (R ampino and Sanders: LOSO: FitzGerald et al., 1994: Schwab et al., 2000). The low mean tidal range (0.7 m) and unlimited fetch offshore Martha’s Vineyard island have combined to produce a high-energy, wave- dominated environment (Figure 1). Furthermore; the existence of a large glacial sand source has been condu- cive to the formation of coastal barriers, which at lower stands of sea level protected muddy coastal bays and fringing saltmarshes (Oldale, 2001). A recent geophysical study of the seafloor offshore Martha’s Vineyard (Goff et al., 2005) offered an opportunity for study of the sub- merged Holocene coastal deposits that contained re- mains of shallow-water macrofauna, including I. obsoleta High-resolution seismic imaging of the seafloor and vibracores, ranging in depth of penetration from 0.5 to approximately 2.0 m, were used to delineate a large sub- merged pe aleo-valley (Figure 1; Buynevich et al., 2002). The main valley (width: 300-500 m: de »pth: 1.5-7.0 m) the offshore extension of Edgartown Great Pond, one of many proglacial spring-sapping valleys incise sd into the late Pleistocene glacial outwash deposits (Uchupi and Oldale, 1994). All of the sediment cores that penetrated marine sands and sampled organic-rich mud facies are confined to the paleo-valley. These deposits contain vari- Page 38 THE NAUTILUS, Vol. 121, No. 1 12 (east valley mean tidal range margin) 13 1.0 ca. 6,200 years ago 44 + 2.0 depth below present MHW (m) 16 seafloor 15 Martha’s Vineyard 6,230+60 calBP 5km paleo-valley aa transect 1com 2.0 eS coarse sand (variable gravel content) eS fine-medium marine sand ee black backbarrier mud with in-situ shells high-marsh peat 1400 2100 3100 3900 19 distance from shoreline (m) Figure 1. Geological section along the axis of the main valley offshore Martha’s Vineyard (see inset for location) showing the occurrence of black, organic-rich backbarrier muds underlying marine sands. These facies contain in-situ gastropods and biv alves and have been sampled to a depth of at least 1S m below present sea level. Photographs: A) Iyanassa obsoleta shells encased in black mud (one shell was used for dating). Saltmarsh peat recovered in the adjacent core 1 has a similar age and both dates are used to constrain the position of mean high water ca. 6,200 years ago. B) cleaned I. obsoleta shell from core 13 (sample depth: ~15 m below present se a level) able amounts of organic material, occasional small bur- constrained portion of the regional sea-level history, but rows, as well as whole shells of I. obsoleta (Figure 1, fit well within the regional sea-level envelope of Oldale photos) and shallow-water bivalves. One core through and O'Hara (1980), as well as New York and New Jersey the valley margin retrieved a short section of saltmarsh shelf data (Stuiver and Daddario, 1963; Rampino and peat, which suggests a low-energy backbarrier setting Sanders, 1980). The similarity between the ages of high- with saltwater access through a tidal inlet. marsh peat and that of eastern mud snail, along with the The age of an in-situ I. obsoleta shell from core 2.4 (14 geological context and ecology of I. obsoleta, demon- m below present mean high water [MHW]) was deter- strates the potential of this species as a paleo-sea-level mined with accelerator mass spectrometry (AMS) radio- indicator. Although the elevation of muddy sediments carbon dating and compared to that of a mee ee peat and macrofaunal remains relative to contemporaneous from core | (13.5 m below MHW). A date of 6,230 + 60 sea level is less constrained than that of high-marsh peat, cal BP (calibrated years before present; 20 error) i‘ the their association with peat and great cross-shore extent shell provides an age for the minimum elevation of mean make them additional points of reference for the position high tide level in this part of the valley fill The age of of sea level. Substantial compaction by loading and vi- 6,245 + 75 cal BP on the saltmarsh peat (8'°C= —15.4%o) bracoring action is unlikely due to relatively small thick- further constrains the paleo-MHW elevation at this lo- ness of overlying sediments and basal positions of dated cation (Figure 1). Both dates plot in a relatively poorly material in both cores, respectively, SSS SSS SSS SS SSS I MV: Buynevich, 2007 Page 39 This study demonstrates that in areas where occur- rence and thickness of peat are limited, such as in mi- crotidal settings, I. obsoleta-bearing muds may prove to be reliable indicators of past water levels, energy condi- tions, and former shoreline positions. In particular, in- tertidal and shallow subtidal portions of submerged val- ley fills provide an ideal geological setting for preserving a nearly continuous sequence of back barrie ccameal that can be used for paleoenvironmental and sea-level reconstruction. ACKNOWLEDGMENTS The study was funded by the Office of Naval Research and the Coastal Ocean Institute of the Woods Hole Oceanographic Institution. Author wishes to thank John Goff, Roy Wilkens, Rob Evans, Peter Traykovski, Chris Jenkins, and the crew of R/V Care HENLOPEN for assis- tance with coring operations and Heidi Fuchs for eco- logical insight. Michael Savarese provided helpful com- ments on the manuscript. LITERATURE CITED Batchelder, C. H. 1915. Migration of Ilyanassa obsoleta, Lit- torina littorea, and Littorina rudis. The Nautilus 29: 43-46. Brenchley, G. A Ilyanassa obsoleta in Barnstable Harbor. Biological Bulle- tin 159: 456-457. Brenchley, G. A. 1982. Predation on encapsulated larvae by adults: effects of introduced species on the gastropod Iya- nassa obsoleta. Marine Ecology Progress Series 9: 255-262. Buynevich, I. V, R. L. Evans, S. Schock, R. Wilkens, J. A. Goff, P. A. Traykovski, C. Jenkins, G. Quentin, P. Beaujean, J. Wulf, C. Vaczo, H. Nieto, and H. Gittings. 2002. Geom- etry and sedimentary characteristics of a submerged pro- glacial spring-sapping valley, offshore Martha’s Vineyard, Massachusetts. Eos Transactions, AGU, 83(47), Fall Meeting Supplement, Abstract OS61A-0156. Carlton, J. T. 1992. Introduced marine and estuarine mollusks of North America: an end-of-the-20'"-century perspective Journal of Shellfish Research 11: 489-505. Culbert. \W. and L. Raleigh. 2001. The Ecology of Coastal Salt Ponds: A Pilot Study at Long Point Wildlife R efuge, West Tisbury and Chilmark, Martha’s Vi ineyard. The Trustees of Reservations, V ineyard Haven, Massi ichusetts, 74 pp. Dexter, R. W. 1961. Mass movement of a colony of mud snails, Ilyanassa. The Nautilus 75: 85-S6. Donnelly, J. P.. P. Cleary, P. Newby, and R. Ettinger. 2004. iG oupling instrumental and geological records of sea-level change: evidence from southern New England of an in- . 1980. Distribution and migratory behavior of crease in the rate of sea-level rise in the late 19"" century. Geophysical Research Letters 31, L05203: 4 pp. FitzGerald, D. M., P. S. Rosen, and S. van Heteren. 1994. New England Barriers. In: Davis, R. A., Jr. (ed.) Geology of Holocene barrier island systems. Springer-Verlag, pp. 3 eae ‘ Goff, J. A., L. Mayer, P. A. Traykovski, I. V. Buynevich, R Wilkens. R. Raymond, G.G lang, R. L. Evans, H. Olson, and ©. Jenkins. 2005. Detailed inve stigation of sorted bed- forms, or “rippled scour depressions”, within the Martha's Vineyard Coastal Observatory, Massachusetts. Continental Shelf Research 25: 461-484. Kelaher, B. P. J. S. Levinton, and J. M. Hoch, 2003. Foraging by the mud snail, Myanassa obsoleta (Say), modulates spa- tial variation in benthic community structure. Journal of Experimental Marine Biology and Ecology 292: 139-157. Oldale, R. N. 2001, Cape Cod, Martha's Vineyard, and Nan- tucket: The Geologic Story, 2"4 Edition, On C Jape Publi- cations, 224 pp. Oldale, R. N. and C. J. O'Hara. 1980. New radiocarbon dates from the inner continental shelf off southeastern Massa- chusetts and local sea-level rise curve for the past 12,000 years. Geology 8: 102-106, Race, M. S. 1982. Competitive displacement and predation between introduced and native mud snails. Oecologia 54: 337-347. Rampino M. R. and J. E. Sanders. 1980. Holocene transgres- sion in south-central Long Island, New York. Journal of Sedimentary Petrology 50: 1063-1080. Robertson, J. R. 1979. fivdeave for tidally correlated feeding rhythms in the eastern mud snail, Myanassa obsoleta. The Nautilus 93: 3540. Stuiver, M. and J. J. Daddario. 1963. Submergence of the New Jersey coast. Science 142: 941. Schwab, W. C., E. R. Thieler, J. R. Allen, D. S. Foster, B. A. Swift, and F. Denny. 2000. Influence of inner-continental shelf geologic frame work on the evolution and behavior of the barrier-island system between Fire Island Inlet and Shin- necock Inlet, Long Island, New York. Journal of Coastal Re- search 16: 408-422. Uchupi, E. and R. N. Oldale. 1994. Spring sapping origin of the enigmatic relict valleys of Cape Cod, Martha’s Vineyard and Nantucket Islands, Massachusetts, Ge omorphology 9: $395. van der Plassche, O. 1986. Introduction. In: Van der Plassche, O. (ed.) Sea-level Research: A Manual for Sample Collec- tion and Evaluation of Sea Level Data. Norwich, Geo Books, 1-26. Weiss, H. M. 1995. Marine animals of Southern New England and New York: Identification keys to common nearshore and shallow water macrofauna. Bulletin 115, State Geo- logical and Natural History Survey of Connecticut, 709 pp. Whitlatch, R. B. 1982. The ecology of New England tidal flats: a community profile. U.S Fish and Wildlife Service, Bio- logical Services Program, Washington, D.C., FWS/PBS- 81/01, 125 pp. THE NAUTILUS 121(1):40, 2007 Page 40 Errata In the last issue of The Nautilus, in the article by Thomas J. DeVries (2006), please substitute the specific epithet meleani for stucchii: On page 139, left-hand column, second paragraph, second line; On page 140, legend of Figure 2, first line; On page 146, left-hand column, sixth paragraph (“Remarks”), first line. Due to an editorial lapse, the word stucchii was used unintentionally in these three instances. LITERATURE CITED DeVries, T. J. 2006. The Neogene history of Prisogaster Mérch, 1850 (Gastropoda: Turbinidae) in South America. The Nautilus 120: 139-149. THE NAUTILUS 121(1):41, 2007 Page 41 Notice THE 2007 R. T. ABBOTT VISITING CURATORSHIP The Bailey-Matthews Shell Museum is pleased to invite applications for the 2007 R. T. Abbott Visiting Curatorship. 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Box 1580 Sanibel, FL 33957 ie er NN oe de Volume 121, Number 2 June 27, 2007 x T 9Q_72 CONTENTS ISSN 0028-1544 Ellen E. Strong On the anatomy and systematics of Juga from western North America Terrence J. Frest (Gastropoda: Cerithioidea: Pleuroceridae) .. 0... ee 43 Timothy A. Pearce Discriminating shells of Gastrocopta pentodon (Say, 1822) and G. Marvin C. Fields tappaniana (C. B, Adams, 1842) (Gastropoda: Pulmonata) with an example Kayoko Kurita from the Delmarva Peninsula, eastern USA... 0. 66 M. G. Harasewych Upper Jurassic Pleurotomariidae (Gastropoda) from Steffen Kiel southwestern Madagascar ... 1... ee 76 M. G. Harasewych Sassia melpangi, a new ranellid species (Gastropoda: Tonnoidea) from Alan G. Beu the Central Pacific 6... ee eee 90 José Carlos N. de Barros A new species of Microcancilla (Gastropoda: Cancellariidae) from the Richard E. Petit continental slope off northeastern Brazil... 0.00. 95 Silvio Felipe B. de Lima A new species of Gerdiella (Gastropoda: Cancellariidae) from the South José Carlos N. de Barros Atlantic Ocean off Brazil with discussion of an undescribed species ...... . 99 Richard E. Petit Book: Review? .(¢ 5.6 4 4 ced oe b4.h4. bs Sameer d ob ew Oe el embed Cwregd e544 G89 phew. ea sede 5 104 Sponsored in part by the State of Florida, Department of State, Division of Cultural Affairs, the Florida Arts Council and the National Endowment for the Arts prts ¢ 5° % \) Qy “ . 2. ae ad a lS ZS a NATIONAL Tp x ENDOWMENT ce) ¥ FOR THE ARTS THE NAUTILUS 121(2):43-65, 2007 Page 43 On the anatomy and systematics of Juga from western North America (Gastropoda: Cerithioidea: Pleuroceridae) Ellen E. Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution P.O. Box 37012 Washington, DC 20013- stronge@si.edu Strong Deixis Consultants 7012 USA Terrence J. Frest 2517 NE 65th Street Seattle, WA 98115 USA tjfrest@earthlink.net ABSTRACT The family Pleuroceridae is a speciose and ecologically impor- tant family of limnic gastropods in North America and eastern Asia. Juga is the only native pleurocerid genus that occurs in Pacific drainage systems of Western North America, but has only recently been accepted as independent from other North American genera and may have affinities to Asian pleurocerids. As such, this genus represents a key piece to the puzzle of pleurocerid systematics. However, published accounts of Juga anatomy are limited to the reproductive system, Consequently, the anatomy of three species is describe d herein: these three taxa represent the type species of the three extant subgenera: Juga (Juga), J. (Calibasis), and J. (Oreobasis). A lectotype is designated for Goniobasis acutifilosa Stearns, 1890, the type species of Calibasis; Melania newberryi, the type species of Oreobasis, is here removed from the synonymy of Juga bulbosa. This analysis confirms that Juga shares many anatomical fea- tures with other North American and Asian pleuroce srids, but is clearly set apart from eastern North American pleurocerids in features of the ovipositor pore, radula, midgut, kidney, and pallial gonoduct. Juga is distinct from all other limnie cerithio- ideans Teno thus far in the form of the midgut crescentic ridge, the configuration of prostate glands, and an evagination of the kidney wall separating the main chamber and bladder. Based on information currently available, unlike molecular data, there is no morphological feature unambiguously linking Juga to Asian pleurocerids. Anatomically, Ore obasis is strikingly similar to Juga sensu stricto and is synonymized with it, whereas Calibasis is retained as a valid taxon. INTRODUCTION The Pleuroceridae Fischer, 1585, is a speciose and eco- logically important family of limnic gastropods occurring in North America and Eastern Asia. Des spite their im- portance. understanding of their systematics is discour- agingly incomplete. The current classification of limnic lineages within the Cerithioidea Fle ming, 1822, is rooted in the works of Thiele (1928, 1929). ake subdivided the heterogeneous Melaniidae Children, 1523 (an invalid name for Thiaridae Gill, IS71) into six subfamilies in- cluding the Pleurocerinae. An alternative classification advanced by Morrison (1954) distinguished only three limnic families: (i) the Pleuroceridae distributed throughout the Americas, Africa, and Asia, (ii) the Mel- anopsidae in Europe, and (iii) the pantropical Thiaridae. While promoting the important notion of several inde- pendent lineages, this concept heavily weighted plesio- morphic (ovipositor and oviparity) and homoplastic (brooding) features and resulted in a highly polyphyletic Pleuroceridae—a view that persisted for over three de- cades (e.g. Ponder and Warén, 1988). Recent work has resolved some of this confusion and supports the distinc- tiveness of many of Thiele’s groupings (e.g. Glaubrecht, 1996, 1999; Ly deard et al., 2002: Kohler and Glaubrecht, 2001, 2003; Strong and Glaubrecht, 2002, 2003; Kohler et al., 2004; von Rintelen and Glaubrecht, 2005). How- ever, molecular data (Lydeard et al., 2002) do not sup- port monophyly of the Pleuroceridae as currently de- fined (e.g. Bouchet and Rocroi, 2005). The analysis of Houbrick (1988) based on morphological data did not include sufficient taxon sampling to adequately assess monophyly of the family. In North America, ple urocerid diversity is highest east of the continental divide where they are represented by seven genera (Athearnia Morrison, 1971, Elimia H. and Adams, 1854, Io Lea, 1831, Leptoxis Rafinesque, 1819, Lithasia Haldeman, 1840, Pleurocera Ratinesque, 1818, and the extinct Gyrotoma Shuttleworth, 1545) and 72 aaa ‘ly 148 species currently considered valid (Johnson et al., 2005). Juga H. and A. Adams, 1554, with 11-12 estimated valid species (Burch, 1989; Turgeon et al., 1998; Johnson et al., 2005), rocerid genus that occurs in the Pacific and Interior drainages from central California to central Washington (Figure 1). Established on the basis of early teleoconch shell sculpture, four subgenera are recognized (Tayloi 1966; Burch, 1989)—three are extant: Juga H. and A is the only native pleu- a] Page 44 THE NAUTILUS, Vol. 121, No. 2 Figure 1. Black line indicates currently Distribution of juga. B known area of contiguous distribution. Black circles represent isolated sites bey ond main distribution. Black squares are loca- tions of extant subgeneric type localities and of material used in this investigation. Adams, 1554 (with plicate early sculpture), Calibasis Taylor, 1966 (with lirate early sculpture), and Oreobasis Taylor, 1966 (with weak to no early sculpture). Idabasis Taylor, 1966 (with plicate and lirate early sculpture) is known only from fossils. Some east Asian species have also been assigned to Juga, but more recently these have been reassigned to Parajuga nate a and Staroboga- tov, 2004 ( Starobogatov et al., 2004): however, this name is unavailable from a hanes standpoint, as no type species was designated. Juga has no discrete conchological feature that distin- guishes it from eastern North American genera and has often been synonymized with Goniobasis Lea, 1862 (a na synonym of Elimia) (e.g. Tryon, 1865, 1873; Pils- bry, 1 899: Walker, 1918: Henderson, 1935a, b: Goodrich, 1942). Taylor (1966) was the first to restore Juga as a valid genus in the modern literature, primarily on the basis of its disjunct biogeographic distribution and the presence of a distinctive ovipositor (see Discussion, be- low). Based on recent molecular (Holznagel and Ly- deard, 2000; Lydeard et al., 2002) and morphological findings (Prozorova and Raschepkina, 2004; Strong, 2005). Juga is supported as distinct from other North \merican pleurocerids and may have ties to those from eastern Asia (see Discussion, below). As such, this genus represents an important and intriguing part of the puzzle of pleurocerid systematics. Thus, the goal of this analysis is to establish the ana- tomical or ganization for the type species of the current extant subgeneric subdivisions within Juga: Juga H. ‘and A. Adams, 1854: type species (by subse- quent designation Baker, 1963) Melania silicula Gould, 1847. Calibasis Taylor, 1966: type species (by original des- ignation) Goniobasis acutifilosa Stearns, 1890. Oreobasis Taylor, 1966: type species (by original des- ignation) Melania newberryi Lea, 1560. This also represents the first comprehensive anatomi- cal investigation of any Juga species; the only information published thus far concerns female reproductive anatomy (Prozorova and Raschepkina, 2004). In the con- text of ongoing morphological and molecular phyloge- netic analyses of pleurocerids and cerithioideans in gen- eral, this information is vitally important for assessing Fe affinities and relationships of taxa currently placed in the Pleuroceridae. MATERIALS AND METHODS Specimens for morphological study were collected by hand or dip net, stored cold until evening, relaxed with menthol in shallow water from the same spring or stream in a broad, open container overnight, and transferred to dilute cold 4-5% formalin the next morning. Specimens were maintained in formalin for 48 hours then trans- ferred to buffered 70% ethanol/10% glycerin/20% water for longer term storage. Voucher wiateral’ is deposited in the National Museum of Natural History in Washington, DC, (USNM) and with Deixis Consultants in Seat- tle, WA. Specimens were examined using a Leica MZ 12,5 bin- ocular microscope with camera lucida: visualization of structures was enhanced with aqueous toluidine blue. Typically 2 to 4 specimens were examined for each organ system investigated, particularly for complex structures (i.e. midgut, nerves), as well as to assess intra-specific and/or cenconel variation in reproductive anatomy. A comprehensive anatomical account is provided for Juga (Juga) silicula—the type species of the genus. Only discrete differences are detailed for J. ( (Calibasis) ) acuti- filosa and J. (Oreobasis) newberryi with comparative re- marks highlighting qualitative differences. As far as can be detenmiied: near ae material was used for this investigation (see details below). A thorough systematic treatment of each species is not provided as a phyloge- netic analysis and comprehensive revision of the genus are forthcoming (Frest et al., unpublished data). Geographic names, road names and numbers, and land ownership data were confirmed using the DeLorme Mapping Washington, Oregon, and Northern California Atlas and Gazetteer, the late »st available USGS 7.5’ series topographic maps, and National Geographic TOPO! 2006. Universal Transverse Mercator (UTM) grid coor- E. E. Strong and T. J. Frest, 2007 Page 45 dinates are based on NAD27 (1927 North American Da- tum). Locality descriptions have been downloaded from Deixis Consultants MolluscDB!'™'. Collector abbrevia- tions are as follows: EJ, Edward J. Johannes; TF, Ter- rence J. Frest. Institution codes cited in the text are: USNM: Na- tional Museum of Natural History, Smithsonian Institu- tion, Washington, DC; MCZ: Museum of Comparative Zoology, Harvard, RESULTS Pleuroceridae Fischer, 1885 Juga (Juga) silicula (Gould, 1847) Type Material: Three syntypes (USNM 12137) (cited as MCZ 12137 in Graf, 2001) (Figures 2-4). Johnson (1964) indicated that the largest (Figure 3) might be the specimen illustrated by Gould ( 1852, 1856, pl 10, figs. 164, 164a); however, in tinct of whorls and overall size, the figured specimen is most likely the smallest syntype (Figure 2). As the figured specimen represents the smallest syntype, and the largest specimen mostly lacks the distinguishing axial ornament characteristic of juvenile shells, a lectotype is not here selected. Type Locality: Indicated as “Nisqually, Oregon” (Gould, 1847) (see Figure 1). These specimens likely ae have been Gollented by the Wilkes Expedition, 1838-1842, before the separation of Washington Terri- tory from the larger Oregon Territory in 1853. This ex- pedition started from Fort Nisqually, which was located in the current Washington State (still part of Oregon Territory in 1547). \. There was no rail station or city with the name “Nisqually” at that time. Thus, the river or the fort would be indicated. Johnson (1964) reported the type locality as “near present site of Tacoma, Washing- ton” (also perpetuated by Graf, 2001), which is loosely true but unnecessarily vague and misleading as Tacoma lies outside the northern range of Juga, w hich ends just north of the Nisqually River (Pierce-Thurston Co. line)—by no mere coincidence near the approximate southem terminus of Late Wisconsinan glaciation. The Nisqually River is a glacial flour stream originat- ing on Mount Rainier with a depauperate fre shwate oT mollusk fauna, despite various literature ascriptions, Its tributaries are comparatively rich, however, especially as they approach Puget Sound. “Nisqually” thus likely re- fers to Fort Nisqually *, one of two trade outposts of the Hudson Bay oa founded in 1833 on the Nisqually Reacl 1 dive -ctly south of Sequalitchew Creek (Hitchman, 1 Puget Sound tributary northeast of the de River. In 1843 the operation was moved about : Dupont, Washington (Phillips, 1997), roughly in the same drainage. For quite some time, this was the only settlement in the area, so that the origin of Gould’s speci- mens could be Sequalitchew Creek or one of the other nearby tributaries of the Nisqually River. Attempts to 2 miles northeast, to approximately Ny present site of collect Juga from the boggy Sequalitchew Creek have been unsuccessful (Frest, unpubl. data). However, ma- terial from nearby McAllister Creek, also a Puget Sound tributary just west of the Nisqually River, has young specimens most closely resembling the types found thus far. Remarks: Evidently a valid species (see also Good- rich, 1942: Burch and Tottenham, 1980: Burch, 1982a, b, 1989; Graf, 2001) but perhaps with a rather restricted range, confined to a few streams on the southeastern end of Puget Sound, near the Nisqually River. Material Examined: Washington: Thurston County: McAllister Creek at eine Road bridge, near McAl- lister Creek Hatchery, depth 0-1.22 m (Zone 10 520890E 5210540N [122°43'34”"E, 47°02'59’N], Nisqually 1994 7.5! quadrangle, elevation 1.5 m), Deixis Consultants locality #5709, 29 Sep. 2005, collected by TF, EJ (USNM 1100657) (9 specimens dissected) (see Figures 7-9). External Anatomy: Operculum ovate, corneous, dark reddish brown in color, with 3.5 whorls; paucispiral with eccentric nucleus of approximately 3 whorls (Figure 16). Nucleus comprising slightly less than one half of total length (~45%). Head- foot dark gray to black in color, with lighter gray snout tip and and pale. foot sole; in females, ovipositor groove pale in color. Foot broad with wide propodium and long anterior pedal gland along anterior margin (Fig- ure 17, ap). Ciliated egg groove forming longitudinally groovy ed tract extending fon anterior tip of palliz il gon- oduct (go) and broadening continuously into shallow: tri- angular shelf on side of neck below right cephalic ten- tacle (ovp). Ovipositor surface groovy ad. with folds di- rected medially into pore. Shallow grooved tract extending from pocket to edge of foot, “short distance back from anterior pedal gland. Extensible snout (Fig- ures 17, 18, sn) broad, squarish, with short triangular cephalic tentacles (t); tentacles also short in relaxed specimens. Ctenidium (Figures 18, 19, 26, et) extending from pos- terior end of mantle cavity to near mantle edge, anteri- orly curving toward the left. Osphradium forming simple ridge alongside efferent branchial vessel, sometimes with anterior tip markedly curving toward the left; osphra- dium undulating slightly at anterior and posterior ends, aan aes (os). Hypobranchial gland moder- ately well developed with transverse ridges, especially at posterior end of mantle cavity (Figures 1S, 19, hg). aca | System: RADULA: Radula comprising ~98 rows (n = 2) (Figures 20-25). Rachidian bro: idly rectan- gular, rr than tall, with smoothly rounded v-shaped lower margin and single small basal denticle at each lower, outer corner (Figure 22). Upper margin slightly concave with cutting edge bearing one large central tri- angular, spatulate cusp, and three stout, conical denticles on each side. Lateral teeth (Figures 20, 21, 23) with moderately short lateral extensions (slightly more than THE NAUTILUS, Vol. 121, No. 2 1S56 pl 10 figs 164. 164a Shells of Juga species. 2-4. Juga (Juga) silicula (USNM 12137, syntypes). Figure 2 is likely the figured specimen uga (Calibasis) acutifilosa (USNM 60596 lectotype Stearns, 1S90 pl 15. fig. 9). 6. Juga (Oreobasis) newberryi (USNM 118961, lectotype). 7-15. Material examined in morphological analysis. 7-9. Juga (Juga) silicula USNM 1100659). 10-12. Juga (Calibasis) acutifilosa (USNM Sc cl lf the length of lateral cutting edge), and single promi- itulate cusp flanked by two inner and two to three nticl Outermost denticle weakly developed nr shape ize, and position (Figure 23 Figures 24, 25) with broadly rounded lon lender shafts. Narrow flanges 1100658). 13-15. Juga (Oreobasis) newberryi (USNM_ 1100660 developed on outer edges of marginal teeth shatts along distal one half to two thirds. Outer flange much wider and longer on inner marginal tooth; outer flange only slightly wider on outer marginal tooth. Inner marginal teeth with six and outer marginal teeth with seven flat tened denticles E. E. Strong and T. J. Frest, 2007 Page 47 ‘int OOO NET ek Or tA UWI 0s-\———— 17 Eee) Figures 16-19. External anatomy of Juga (Juga) silicula (USNM 1100659). 16. Operculum. 17. Ovipositor and egg groove. Right lateral view of head-foot. 18-19. External view of organs in visceral mass. Dotted line indicates extent of pericardium under main kidney chamber (Figure 15). Abbreviations: ag, albumen gland; ap, anterior pedal gland; b, bladder; eg, capsule gland; em, columellar muscle: et, ctenidium; dg, digestive gland; f, foot; go, pallial gonoduct; hg, hypobranchial gland; int, intestine; kd, main kidney chamber; me, mantle edge; op, operculum: os, osphradium; ov, ovary; ovp, ovipositor; per, pericardium; sn, snout; ss, style sac: sto, stomach: t, cephalic tentacle. Scale bars = 1 mm. Page 45 THE NAUTILUS, Vol. 121, No. 2 Figures 20-25. Radula of Juga (Juga) silicula (\USNM 1100659). 20. Section of anterior radular ribbon. Scale bar = 200 jm. 21. Rachidian and lateral teeth. Scale bar = 50 pm. 22. Detail of rachidian teeth. Scale bar = 20 pm. 23. Detail of lateral teeth; note weakly formed outer third denticle. Scale bar = 50 zm. 24. Marginal teeth. Scale bar = 50 wm. 25. Detail of cutting edge of marginal teeth. Note unequal size of cusps on inner and outer teeth. Scale bar = 20 jm. Forecut: Buccal mass short and robust (Figure 26, bm). Odontophore occupying posterior one half to two thirds of buccal cavity with small, glandular subradular organ protruding before radula. Small jaws present at anterior ends of dorsal folds; epithelium of buccal cavity between dorsal folds glandular (stippled). Dorsal folds deeply cleft along midline adjacent to odontophore; cleft receives salivary gland ducts at posterior end and shallows ante- riorly. Very shallow, non- -glandular buccal pouches ex- page underneath Bosal folds adjacent to buccal gan- lia (bg) at rear of buccal cavity. Radular sac (1s) shit a 4.4 mm), curving upward behind base of buccal mass, not passing through nerve ring, with tip resting against right posterior Sad of buccal mass. Robust buéeal retrac- tors (rt) inserting onto lateral walls of cephalic hemocoel anterior to nerve ring. Short, glandular mid-ventral fold forming small triangular ridge just behind odontophore in anterior esophagus, fiesled by two ventro-lateral folds. Ventro-lateral folds converging short distance behind mid- ventral fold, forming rear-facing triangular depression sur- rounding mid-ventral fold. Paired dorsal and ventral folds continuing through long mid-esophagus (e) into posterior esophagus. Epithelium of mid-esophagus between dorsal and ventral folds longitudinally grooved; septate esophageal gland lacking. Posterior esophagus narrow, with numerous folds of approximately equal height. Long, tubular salivary glands (sg) passing through circum- esophageal nerve ring, extending . posterior esophagus. Mipcut: Esophagus opening under ledge on left side of midgut floor (Figure 27, e). Marginal fold (mf) extending anteriorly from esophageal aperture alongside major typhlosole (t1), then turing posteriorly, bordering right margin of sorting area (Sa). Sorting area elongate, rect- angular, tapering posteriorly; posterior tip curving slightly to the left around crescent-shaped sorting area pad (sap). Accessory marginal fold (amf) forming. weak ridge paralleling marginal fold from near esophagus, curving around posterior margin of sorting area; poste- rior segment of fold variable, in some specimens inter- secting straight longitudinal ridge along left posterior end of sorting area (figured), in other specimens weakly bifurcate (see e.g _ Figures s 37, 51). Fine parallel striations extending anteriorly from esophagus up face of major typhlosole (Figure 27, tL). Midgut roof to the left of sorting area coarsely folded and cuticularized (eu). Gas- tric shield (gs) broadly concave; shield continuous with cuticle of adjacent oe of stomach roof and floor. Glandular pad (gp) large, rounded posteriorly, with lightly textured surface. Slight ov i: lip of glan- delar pad forming shi low pocket ( y behind gastric shield. Crescentic. ridge (er) ee wide, shallow crescentic groove. Proximal end of ridge posteriorly bor- dering deep pouch that receives multiple openings of digestive gland (dd); distally, ridge fusing to right, pos- terior end of glandular pad. Size of opening to digestive eland duct ve stibue variable. Single, weak, irregular lon- citudinal fold (ef) along floor opposite caecum. Promi- nent ioe fold (uw) extending from beneath right side of style sac lip (ss), along flog. to ridge extending from base of major ty phlosole: fold bounding u- -shaped depression below lip of style sac. Style sac large, com- municating along entire length with intestinal eroove; intestine forming prominent protuberance at distal tip of E. E. Strong and T. J. Frest, 2007 Page 49 26 Figures 26-29. Intemal anatomy of Juga (Juga) silicula (USNM 1100659). 26. Mantle cavity and anatomy of cephalic hemocoel. Dorsal view, anterior is below. 27. Midgut anatomy. Dorsal view, anterior is uppermost. 28. Kidney anatomy. Internal view of bladder. Lateral view, anterior is uppermost. Roof of bladder (below intestine) cut open to reveal interior; adjacent to incision, stippling indicates intersection of Ory tubules with roof of bladder. Arrow indicates opening in outpocketing of wall between bladder and main kidney chamber. 29. Circum-esophageal nerve ring. Frontal view on the left, right lateral view on the right. Arrow indicates connective to supra-esophageal ganglion. Abbreviations: amf, accessory marginal fold; arv, afferent renal vessel: bg, buccal ganglion: ec, caecum: ce, cerebral ganglion: cf, caecal fold: co, thickened connective between left pleural and sub-esophage sal vaneglia; cr, crescentic ridge; et, ctenidium; cu, cuticularized region of stomach roof; d, dialyneury; dd, digestive gland duct vesuivule: e, esophagus; go, pallial gonoduct; gp, glandular pad; gs, gastric shield; int, intestine; kd, main kidney chamber: me, mantle edge; mf, marginal fold: np, nephropore: os, osphradium; pe, pedal ganglion; pl, pleural ganglion; r, rectum; rs, radular sac; rt, buccal retractor muscle; sa, sorting area; sap, sorting area pad; sb, sub- -esophageal ganglion; sg, salivary gland; sp, supra-esophageal ganglion: ss, lip of style sac: st, statocyst: t, cephalic tentacle; t1, major typhlosole; u, u-shaped fold; z, zygoneury. Scale bars = 1 mm Page 50 THE NAUTILUS, Vol. 121, No. 2 style sac where it separates from the latter (not visible dorsally; see e.g. Figure 50, ss). Crystalline style present. HINDGUT: Proximal intestine (Figures 18, 19, int) pass- ing below distal tip of style sac, then extending posteri- orly in broad loop, partially overlying style sac, to main gastric chamber (sto). Intestine continuing forward, passing under posterior end of main indaey chamber (kd), entering pallial roof between bladder (Figure 19, b) and main kidney chamber (kd), to papillate anus near mantle margin (Figure 26, r). Reno-pericardial System: Kidney comprising two interconnected chambers (Figure ‘ 28). Main chamber (kd) lying along dorsal surface of body whorl, anteriorly surrounding pericardium (Figure 18, per, dotted line), crossing axis of body from right to left and extending short distance into pallial ry at base of mantle cavity. Main chamber occluded with excretory tubules anteri- orly (within pallial roof), posteriorly, and along left mar- gin. Central portion of main chamber with small, narrow Tannen Second chamber (exposed chamber in Figure 28) extending between pericardial chamber to right body wall below intestine, forming small bladder (Figure 19, b). Wall separating main chamber and bladder forming large outpocketing; small aperture within wall, just in front of afferent renal vessel, connecting main chamber and bladder (Figure 28, arrow). Bladder io gely occluded by vertical sheets of excretory tissue radiating from af- ferent renal vessel (arv). Sheets of excretory tissue branching and anastomosing, and fusing to right lateral wall, floor and roof below intestine; sheets laterally en- closing outpocketing of main chamber wall. Bladder communicating to mantle cavity via large nephropore (np); outpoc ke ting of main chamber wall exte nding into ne Bee restricting communication with m: antle ¢ cav- ity. Bladder penetrating connective tissue along right ee of body, short distance into mantle cavity. Nephridial gland absent. Pericardium long and narrow (Figure 1S, per, dotted line), extending to recurved intestinal loop. Nervous System: Circum-esophageal nerve ring (Figure 29) lying immediately behind buccal mass, well behind base of cephalic tentacles. Cerebral ganglia (ce) connected by short, stout commissure, each producing seven nerves (optic, statocyst, tentacular, and 4 labial nerves). Buccal ganglia (Figure 26, bg) lying ventro- laterally at base of buccal mass, adjacent to retractor muscles, at outer edges of esophagus where it emerges from buccal cavity. Pleural ganglia (Figure 29, pl) ) lying behind and below cerebral vanglia, connected to cere- bral ganglia by short, thick connectives. Pedal ganglia pe) with two prominent anterior nerves and variable number of smaller accessory nerves (five to seven). Sta- toeysts (st) with numerous statoconia present dorsally alongside pedal ganglia behind pedal connectives, Sub- esophageal ganglion (sb) joined to left pleural canglio by thickened connective (co); connective producing on small nerve. In addition to connectives to right and left pleural and visceral ganglia, sub-esophageal ganglion producing four small nerves. Zygoneury (z) formed be- tween sub-esophageal and right pleural ganglia. Long connective uniting right pleural and supra- esophageal ganglia (Figures 26, 29, sp), the latter lying on left side of mentie floor near midline of osphradium. Dialyneury formed between pallial nerve of left pleural ganglion and osphradial nerve of supra-oesophageal ganglion at junc- tion of mantle roof and floor. Single visceral ganglion present between pericardium and kidney at base of mantle cavity. Reproductive System: FEMALE: Gonad (Figures 1S, 19, ov) dorsally surrounding digestive gland from tip of visceral mass to posterior end of midgut (sto). Oviduct emerging ventrally from ovary. Renal oviduct (Figure 30, ovi) entering glandular pe allial oviduct at base of mantle cavity. Palliat oviduct with proximal albumen (ag) and distal capsule glands ( (cg). Proximal albumen gland, be- low pallial portion of bladder, forming u- shaped tube with glands developed dorsally along axis of fold and simple gonoductal groove between opposing flattened surfaces of glands. Anteriorly, albumen and capsule glands becoming highly glandular and thrown into com- plex undulating folds: gonoductal groove compressed and highly eonvohite d (gg). Albumen gland approxi- mately one third the length of capsule | land. Pallial ovi- duct communicating with mantle cavity through narrow aperture along entire length (arrow), except for tubular section of alleunien gland at base of mantle cavity. Above aperture, along anterior ~one fourth of f oviduct, deep sperm gutter (sg, dashed line) present within medial lamina; gutter opening broadly to long, broadly rounded spermatophore bursa ( (spb). Bursa broadening posteri- orly and extending to tubular portion of aibeanen gland. Behind opening to bursa, sperm gutter becoming shallow abruptly and continuing posteriorly ( (dashed line) as shal- low groove. Near posterior end of oviduct, shallow ridge entering small rounded aperture, just inside ventral edge of eden lamina, leading to small pouch-like seminal receptacle (res). Prominent glandular protuberance from opposing inner surface of lateral lamina extending into receptacle aperture, completely filling narrow proxi- mal portion. Thin, narrow glandular ridge extending an- teriorly from protuberance along inner edge of lateral lamina; glandular ridge extending to pallial oviduct tip just inside seminal groove. Mate: Narrow vas deferens (Figure 31, vd) emerging ventrally from testes, continuing forward along ventral midline of whorl. Short distal portion of vas deferens thickened and forming straight seminal vesicle. Vas det- erens narrowing and curving dorsally to enter posterior end of prostate (pr) at base of mantle c ‘avity. Prostate glandular, opening to mantle cavity through narrow ap- erture along entire length except for a short fused seg- ment at base of mantle cavity (Figures 31, 33, arrow). Glands of medial and lateral laminae closely appressed, forming tightly interlocking tongue and groove arrange- E. E. Strong and T. J. Frest, 2007 Page 5] Figures 30-33. Reproductive anatomy of Juga (Juga) silicula (L Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. “< indicates transition between albumen and capsule glands. 31. External, left lateral view of prostate. Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. 32. Internal view of prostate medial lamina. Anterior is to the right. 33. Internal view of prostate lateral lamina. Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. Abbreviations: ag, albumen gland; eg, capsule gland: gg, gonoductal groove; ovi, renal oviduct; pr, prostate; res, seminal receptacle; sg, sperm gutter; spb, spe rmatophore bursa; vd, vas deferens. Scale bars = 1 mm. ment (Figures 32, 33). Glandular tissue of medial os ventrally forming elongate, flattened textured fold (Fig- ure 32), corresponding to concave a surface of op- posing fold in lateral lamina (Figure 33). Medial lamina expanding to surround fold within fe ral lamina. Dor- sally and posteriorly, glands of lateral and medial laminae flaring to form flattened flange; size and shape of flange variable between individuals. Apart from fold in medal lamina, inner surface of prostate essentially smooth. Juga (Calibasis) acutifilosa (Stearns, 1890) Type Material: Lectotype (USNM 60596; figured specimen in Stearns, 1890, pl. 15, fig. 9) (Figure 5), by present designation, in order to e ananee the stability of the nomenclature in accordance with article 74.7.3 of the ISNM 1100659), 30. External, left lateral view of pallial oviduct. ICZN; indicated as holotype on label. Fourteen paralec- totypes (USNM 60596X) in secondary type collection of USNM, indicated as paratypes on old Jabel and as syn- types on newer label printed in 2001. Although Graf (2001) stated that the “holotype” is by original de ssigna- tion, Stearns figured the largest of the syntypes and pro- vided the dimensions, but did not make an explicit : pe designation (designation of holotype) in the text and in- dicated that the description was based on examination of approximately three dozen specimens. Collected by H. W. Henshaw. Type Locality: “Eagle Lake” (Stearns, 1890). Taylor (1981) corrected this to head of Willow Creek, Lassen Co., California (see Figure 1). There are no populations in the Eagle Lake drainage which includes several small THE NAUTILUS, Vol. 121, No. 2 tributaries that flow into the lake during winter and the only outflow is through the remnants of ihe Bly Tunnel. Initiated during the 1920’ s, the Bly irrigation project di- verted water from Eagle Lake to Willow Creek at Mur- rers Meadows sev eal kilometers away in the more arid Honey Lake drainage. Today, the ‘tunnel is mostly blocked except for an 8’(~20 cm) pipe that still allows some outflow to Willow Creek. The headwaters of Wil- low Creek are regarded as springs along Murrer’s Upper Meadow and Bly Tunnel (Moyle et al., 1996). Graf (2001) lists the type locality as “Eagle Lake, [Lassen Co.,] California’. Remarks: The species varies widely in shell morphol- ogy, from populations with several strong persistent lirae to those with lirae confined to the adapical whorls. Body color also may differ drastically from population to popu- lation. Few sites show much intrapopulation variance; but adults can vary from population to population, with those at some sites highly variable and others essentially invariant. Very few populations i in onlya part of the range are as strongly and completely lirate as some Willow Creek (and Murrer’s Upper Meadow) populations. These observations are consistent with preliminary re- sults based on COI sequences that indicate the species, as currently recognized, is highly polyphyletic (Frest et al., unpublished data). Material Examined: California: Lassen County: ie low Creek collected off dirt road (to E.), 0.40 kin S. Murrer’s Lower Meadow, depth 0.05-0.20 m (Zone if 695000E 4493820N [120°41'51"E, 40°34'26"N], Gall- atin Peak 1989 7.5’ quadrangle, elevation 1509 m), Deixis Consultants locality #1484, 10 Sep. 1993, col- lected by TF, EJ] (USNM 1100658) (4 specimens dis- sected) (see Figures 10-12); southern-most of three springs at N. end of ae Lower Meadow, below road on E. side of meadow, E. of Eagle Lake, depth 0-0.03 m (Zone 10 694995E 4495225N [120°41'49”E, 40°35'12"N], Gallatin Peak 1989 7.5’ quadrangle, eleva- tion 1545 m), Deixis Counsultants locality #1287, 10 Sep. 1993, collected by TF, E] (USNM 1100659) (3 speci- mens dissected). External Anatomy: Operculum ovate, with slightly angular tip (Figure 34). Nucleus comprising slightly less than one half of total length (~41%). Ovipositor pore rather deep (Figure 35, ovp) with deep, highly grooved tract extending to edge of foot, slight doa. back from anterior pedal gland ( (ap). Mantle edge (Figure 35, me) cre mulated, cor respond- ing to spiral teleoconch sculpture. Ctenidium (Figure 36, ct) extending from posterior end of mantle cavity to mantle edge. Hypobr: mchial gland weakly developed ( (hg). {EMARKS: With the exception of minor individual and/or preservational differences, the external anatomy is essen- tially identical to Juga silicula. Juga acutifilosa differs only in that the operculum is slightly more angular and the nucleus ¢ omprise Si a slightly smé le or proportion of the total length, the ovipositor and groove to the « dge of the foot are significantly deeper and more grooved, the groove opens to the foot nearer the end oF the anterior pedal ¢ gland, the gill extends slightly nearer the mantle edge, and the hypobranchial g gland j is much more weakly developed. Alimentary System: RaApuLA: Radula comprising ~104 rows (n = 2) (Figures 38-43). Rachidian basal mar- gin concave bordering bluntly rounded median projec- tion: basal denticles feline or only slightly developed (Figures 39, 40). Cutting edge bearing one lar ge central conical cusp, and two stout, conical dente les on each side (Figure 40). Lateral teeth (Figures 38, 39, 41) with short lateral extensions (slightly less than half the length of lateral cutting edge), and single, prominent triangular cusp flanked by two inner triangular denticles and three to four outer denticles. Wealdy developed outermost denticle may be present or absent (Figure 41). Marginal teeth (Figures 42, 43) with broadly rounded cutting edges ae long, slender shafts. Inner marginal teeth wai five and outer marginal teeth with six fletiened denticles. Forecut: Radular sac long (~6.7 mm), extending back through nerve ring approximately one. half distanee to supra- “esophageal ganglion, then curving anteriorly with tip overlying nerve ring. Long, thin tubular salivary glands passing through circum- esophageal nerve ring, ex- tending to posterior “esophagus. Mine uT: Sorting area elongately triangular (Figure 37, Sa). Accessory marginal fold (amf) forming cereal ridge eee mar ginal fold from near esophagus, curving around posterior margin of sorting area, with weakly be furcate posterior sad Glandular pad (gp) ) moderately large, rounded else with coarsely textured sur- face. Deep pocket (c¢) extending under elendular pad behind gastric shield. Crescentic ridge (er) bounding narrow, shallow crescentic groove. Sty le sac ee intes- tine forming slight protuberance at distal tip of style sac where it separates from the latter (Figure 36, ss). REMARKS: In eae to Juga silicula, the denticles of the rachidian and lateral teeth are typically more conical, with fewer denticles present on the rachidian and mar- ginals, but more outer denticles present on the lateral teeth. The weakly developed outermost denticle on the lateral teeth is variably developed, and only very small. Rachidian basal denticles are lac ‘king, or only sli ghtly de- veloped. The radular sac is exc eptionally | ong in this spe- cies and the salivary glands appear narrower. Within the midgut, in spite of the differences high- lighted above, overall configuration and proportions of features are very similar be tween fuga silicula and J. acutifilosa, with the exception that the sorting area is more elongately triangular in shape in J. acutifilosa. No significant differences in the configuration of the hindgut, with the possible exception that the hindgut dramatically widens upon entering the mantle cavity in the specimens examined for Juga acutifilosa. However, this may be individual variation. E. E. Strong and T. J. Frest, 2007 Page 53 Figures 34-37. Anatomy of Juga (Calibasis) acutifilosa (USNM 1100659, except when noted), 34. Operculum. 35. Ovipositor and egg groove (USNM 1100638). Right lateral view of head-foot. 36. External view of organs in visceral mass (USNM 1100658). Dotted line indicates extent of pericardium under main kidney chamber. 37. Midgut ani itomy. Dorsal view, anterior is uppermost. Abbre- viations: amf, accessory marginal fold: ap, anterior pedal ; gland: e, caecum: ef, caece il fold: eg, capsule gland; er, crescentic ridge: ct, ctenidium; cu, cuticularized region of stomach roof; dd, digestive gland duct vestibule; dg, digestive gland; e, esophageal aperture; f, foot: gp, glandular pad; gs, gastric shield: hg, hypobranchial gland; int, intestine; kd, main kidney chambe r; me, mantle edge: mf, marginal fold: op, pee eh os, osphradium; ov, Ovary; Ovp, ovipositor; sa, sorting area; sap, sorting area pad; sn, snout: ss, style sac: sto, stomach; t, cephalic tentacle; t1, major typhlosole; u, u-shaped fold; v, ventricle. Scale bars = 1 mm Page 54 THE NAUTILUS, Vol. 121, No. 2 Figures 38-43. Radula of Juga (Calibasis) acutifilosa (USNM 1100659). 38. Section of anterior radular ribbon. Scale bar = 100 um. 39. Rachidian and lateral teeth. Scale bar = 50 jm. 40. Detail of rachidian teeth. Scale bar = 50 pm. 41. Detail of lateral teeth. Note very weak yale tes of tiny outer fourth denticle. Scale bar = 50 jum, 42. Marginal teeth. Scale bar = 20 ym. 43. Detail of cutting edge of marginal teeth. Note unequal size of cusps on inner and outer teeth. ‘Seale bar = 20 wm. Reno-pericardial System: REMARKS: Configuration of features within the kidney and density of excretory tissue essentially identical to Juga silicula, with the only exception being that the central lumen within the main chamber is slightly shorter. Nervous System: Pedal ganglia (pe) with two promi- nent anterior nerves and four smaller accessory nerves. REMARKS: Configuration of nervous system, including number of nerves produced by major ganglia, otherwise identical to Juga silicula. ey ane System: FEMALE: Proximal albumen gland (Figure 44, ag), below pallial portion of bladder, forming small, flattened rounded pouch with glands de- veloped along dorsal axis: shape of pouch somewhat vari- able. Along anterior ~one third of oviduct, es sperm gutter (sg, dashed line) present within medial lamina: gutter opening broadly to long, broadly rounded sper- matophore bursa (spb). Behind opening to bursa, sperm gutter shallowing abruptly to shallow groove (dashed line), then rapidly becoming obsolete. Near posterior end of oviduct, small rounde ‘d aperture, just inside ven- tral edge of medial ee leading to small narrow semi- nal receptacle (res). Low, flatly rounded glandular pro- tuberance from oa inner surface of lateral lamina slightly extending into rece oS aperture. Extremely thin, ol undular ridge extending from protuberance to ovi- duct anterior tip along inner “edge of lateral lamina op- posite seminal groove; ridge thickening somewhat ante- riorly. MALE: Glands of medial and lateral laminae forming loosely interlocking tongue and groove arrangement (Figures 46, 47). REMARKS: In contrast to Juga silicula, the proximal albu- men gland forms a small, flattened, rounded pouch. The height and shape of the pouch is variable in J. acutifilosa, but is distinctly smaller and slightly less glandular than the more u- shaped tube in J. cilia ula. Ov exall. the capsule and albumen glands are similar in proportion, but the albumen gland is slightly shorter and not as massive as that in J. silicula. In addition, the seminal groove is longer and deeper anteriorly but becomes obsolete be- fore reaching the seminal receptacle, the lateral lamina glandular ridge is much weaker and the protuberance into the aperture of the seminal receptacle is more bluntly rounded than in J. silicula. Male reproductive anatomy is very similar to that of Juga silicula in the morphology of the folds within the prostate, but the folds are not as highly developed and hence, do not tightly interlock to the same degree. In addition, the ventral fold of the medial ian: is less textured and the dorsal and posterior flange is narrower in ]. acutifilosa. Juga (Oreobasis) newberryi (Lea, 1860) Type Material: Lectotype designated by Graf (2001) (USNM 118961; figured specimen in Lea, 1863, pl. 37, fig. 135) (Figure 6); indicated as holotype on label. Twalvé parale sctotypes ( (USNM LIS9GLX) in secondary type collection of USNM; indicated as paratypes on old label and as syntypes on newer label printed in 2001. At the time of publication, the lectotype designation of Graf E. E. Strong and T. J. Frest, 2007 Page 55 GL rate erik: Zo ae SIS 0 A cE Figures 44-47. Reproductive anatomy of Juga (Calibasis) acutifilosa (USNM 1100659), 44. External, left lateral view of pallial oviduct. Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. “<” indicates transition between albumen and capsule glands. 45. External, left lateral view of prostate. Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. 46. Internal view of prostate medial lamina. Anterior is to the right. 47. Internal view of prostate lateral lamina. Anterior is to the left. Abbreviations: ag, albumen gland; eg, capsule gland: gg, gonoductal groove; ovi, renal oviduct; pr, prostate; res, seminal receptacle; sg, sperm gutter; spb, spermatophore bursa; vd, vas deferens. Scale bars = 1 mm. was invalid as it did not follow strict guidelines concern- ing the language of lectotype designations after 1999, However, under Declaration 44, Amendment of Article 74.7.3 of the ree Bulletin of Zoological Nomenclature 60(4) December 2003), Graf's lectotype designation is now valid. Collected by J. S. Newberry. Type Locality: Indicated as “Upper des Chutes River, Oregon Territory” (Lea, 1560) (see Figure 1). Most likely, the lectotype came from the Deschutes R. near Bend, Oregon, even though pleurocerids are now absent this far up the river. However, Juga newberryi does occur in the lower Deschutes River. Remarks: Lea’s (1860, 1562, 1563) description of Mela- nia newberryi and 1863 illustration are quite accurate. As noted and illustrated by Lea, the most frequent color pattern is three yellow bands separated by three almost black. Occasionally, the lower band may be divided into two or more (Burch, 1989: fig. 452) or the dark bands may be tan in color. Alternatively, the shell may be band- less, in which case the color varies from yellowish-tan to dark tan. Lea (1863: 301) does not seem to have had any of the band-less form; recent field surveys have not re- vealed any “pure” populations of the band-less form, on the other hand, large popul itions are not likely to lack it (Frest, unpublishe .d data) This species is one of a small group of Juga with the whorls smooth throughout ontogeny which led Taylor (1966) to make it the type of Oreobasis. In the same Page 56 THE NAUTILUS, Vol. 121, No. 2 publication, 7 Taylor suggested that J. newberryi is a “probable” synonym of Melania bulbosa Gould, 1847. By 1977, Taylor (1977) had accepted this synonymy without qualification, which was followed by many authors (Burch and Tottenham, 1980; Burch, 1982a, b, 1989: Graf, 2001). However, it is not clear why the two species were considered so ee Comparison of the types in- dicates that, while being similar in whorl height, ‘the two differ in shape of the aperture, whorl profile and rate of whorl expansion, particularly for the body whorl; J. bul- bosa generally has more than three not corroded whorls while J newberryi is often more severely corroded but the early teleoconch of the latter is quite distinctive when present. A search of major museum collections failed to locate many specimens aside from the types; most mu- seum lots ascribed to J. bulbosa or J. newberryi clearly do not belong to either. Tryon (1865) was apparently the first to reflect on the similarity of the two, indicating the presence of bands in Juga newberryi as the sole separating feature, but he did not synonymize them. However, Tryon’s illustration of J. bulbosa (e.g. 1873, fig. 496) seems to use his own speci- men rather than Gould's types, even though he claimed to have had them (1873: 255). In contrast to Gould's types, the specimen figured by Tryon is rather large, strongly corroded with Tes than 3 whorls, and the sur- viving whorls appear strongly convex, with a very deep suture. It is likely a band-less form of J. newberryi. Thus, although long considered synonyms, the pe reception that the two are “exactly similar in outline” ( Tryon, 1865: 246) may be due to Tryon’s apparent confusion between J. bulbosa and bandless forms of J. newberryi. The scarcity of museum lots may also have contributed to the confu- sion about the morphology and occurrence of both taxa. Consequently, we here remove Juga newberryi from the synonymy of J. bulbosa. At present, J. newberryi occurs only in the lower Deschutes River, Oregon, from about Troutdale to roughly 6 miles above the mouth, where it is replaced by J. (Juga) plicifera. Juga bulbosa is likely also a valid species, but attempts to recollect this taxon anywhe re in the historically identified range have been unsuccessful (Frest, un published data): thus, this hypothesis cannot be tested 7 the moment. Material Examined: Oregon: Jefferson County: Des- chutes River at RM 97.2-97.4 on E side of channel, ~0.3-0.6 km S of US 26 bridge and mouth of Shitike Creek, at Rainbow Landing, depth 0-0.41 m (Zone 10 640460E 4957320N M21°13'36"E. 44°45'26"N], Eagle Butte 1962 7.5’ quadrangle, elevation 440 m), Deixis Consultants locality #2472, 13 Aug. 2000, collected by F, EJ (USNM 1100660) ( (10 specimens dissected) (see Figures 13-15). External Anatomy: Operculum ovate, with angular tip (Figure 48). Nuc leus comprising about one third of total length (~34%). Rather shallow ovipositor pore (Figure 49, ovp) with shallow, grooved tract extending to edge of foot, slight distance back from anterior pedal oland (ap). —— Ctenidium (Figure 50, et) extending from posterior end of mantle cavity to mantle edge. Hypobranchial gland (hg) moderate sly well dev oe with warty tex- ture, particularly at posterior end of mantle cavity. Remarks: The external anatomy of Juga newberryi is almost identical to the two preceding species, and differs in that the operculum is more angular in shape and the nucleus is considerably smaller and more basal than ec- centric; as in J. acutifilosa, the gill extends slightly nearer the mantle edge and the ovipositor groove opens nearer the end of ie anterior pedal gland, but the pore and distal groove are considerably “deeper in J. acutifilosa than in ihe other two species. The hypobranchial gland of J. newberryi is unique in having a warty texture. Alimentary System: RADULA: Radula comprising ~106 rows (n = 2) (Figures 52-57). Rachidian with v-shaped lower margin; basal denticles lacking (Figures 53, 54). Cutting edge bearing one large central conical cusp, and two stout conical denticles on each side (Fig- ure 54). Lateral teeth (Figures 52, 53, 55) with short lateral extensions (slightly ee than half the length of lateral cutting edge), and single, prominent triangular cusp flanked by three to four inner and three to four outer denticles. Outermost denticle present, weakly de- veloped, and variable in shape, size, and position (Figure 55). Marginal teeth (Figures 56, 57) with broadly rounded cutting ne and long, slender shafts. Inner marginal teeth with four to five and outer marginal teeth with six to seven flattened denticles. ForEGUT: Radular sac moderately long (~6.25 mm), ex- tending back through nerve ring, then curving upward be baie base of buccal mass with tip overlying left side of nerve ring, Epithelium of mid-esophagus ‘between dorsal and ventral folds irregularly te xtured. Mipcut: Sorting area short, broadly triangular (Fig 51, sa). Accessory marginal fold (amf) forming weak ridge par alls ling marging i fold from near esophagus, curving around posterior me ein of sorting area, with weakly te furcate posterior end. Glandular pad (gp) moderately large, rounded ae riorly, with lightly textured surface. Crescentic ridge (er) bounding shi allow, broad crescentic groove, Style sac small (Figure 50, ss); intestine forming prominent protuberance at distal tip of style sac where it separates from the latter. ReMARKS: As in Juga acutifilosa, the denticles of the rachidian and lateral teeth are typically more conical in J. newberryi than in J. silicula; there are only two outer denticles on each side of the central rachidan cusp, more outer denticles on the lateral teeth, the rachidian basal denticles are lacking, and the lateral extensions are pro- portionally smaller. However, in contrast to J. acutifilosa, J. newberryi and J. silicula share a similar lower margin on the rachidian and similar development of the weak, outermost denticle on the lateral teeth. Juga newberryi is unique in possessing three to four inner denticles on the E. E. Strong and T. J. Frest, 2007 Page 57 Figures 48-51. Anatomy of Juga (Oreobasis) newberryi (USNM 1100660). 48. Operculum. 49. Ovipositor and egg groove. Right lateral view of head-foot. 50. External view of organs in visceral mass. Dotted line indicates extent of pericardium under main kidney chamber. 51. Midgut anatomy. Dorsal view, anterior is uppermost. Abbreviations: amf, accessory marginal fold; ap, anterior pedal gland: au, auricle: b, bladder; e, caecum; ef, caecal fold; er, crescentic ridge; et, ctenidium; eu, cuticularized region of stomach roof: dd, digestive gland duct vestibule: dg, digestive gland; e, esophageal aperture; f, foot: gp, glandular pad; gs, gastric shield; hg, hypobranchial gland: int, intestine; kd, main kidney chamber; me, mantle edge; mf, marginal fold; op, operculum; os, osphradium; ovp, ovipositor: sa, sorting area; sap, sorting area pad; sn, snout; ss, style sac; sto, stomach; t, cephalic tentacle; te; testes; tL, major typhlosole: u, u-shaped fold. Scale bars = 1 mm Page 58 THE NAUTILUS, Vol. 121, No. 2 TRIN — Figures 52-57. Radula of Juga (Oreobasis) newberryi (USNM 1100660). 52. Section of anterior radular ribbon. Scale bar = 100 pm. 53. Rachidian and lateral teeth. Scale bar = 50 pm. 54, Detail of rachidian teeth. Scale bar = 20 pm. 55. Detail of lateral teeth; note weakly formed outer fourth denticle. Scale bar = 50 jum. 56. Marginal teeth. Scale bar = 50 wm, 57. Detail of cutting edge of marginal teeth. Note unequal size of cusps on inner and outer teeth. Scale bar = 20 jm. lateral teeth. The radular sac is intermediate in length relative to the buccal mass compared to J. silicula with a very short sac, and J. acutifilosa with a very long sac. Juga newberryi differs greatly from the two preceding species in the relative proportions of features in the mid- gut, which is significantly longitudinally compressed. Consequently, the sorting area is broadly triangular rather than elongate and thé glandular pad is dispropor- tionately smaller. Possibly reflecting the smaller size of the style sac, the opening of the style sac to the stomach is also consistently smaller. There are no significant differences in the configura- tion of the hindgut between Juga newberryi and J. sili- cula. Reno-pericardial System: REMARKS: In contrast to both preceding species, the bladder of Juga newberryi is compressed and less voluminous, bringing the nephro- pore closer to the afferent renal vecel and decreasing the length of the wall between the main chamber and fhe bladder. Additionally, the sheets of excretory tissue in the bladder are more numerous, and more highly branched, almost entirely occluding the lumen. Nervous System: Pedal ganglia with two prominent anterior nerves andl five to six smaller accessory nerves. In addition to comnections to right and left pleural and visceral ganglia, sub-esophageal ganglion producing three small nerves. {EMARKS: Apart from the differences highlighted above, configuration of the nervous system is basically identical to Juga silicula. Reproductive System: FEMALE: Capsule gland (Fig- ure 58, eg) comprising approximately anterior two thirds of pallial oviduct. Mong anterior ~one third of oviduct, shallow sperm gutter (sg, dashed line) present within medial lamina; gutter opening narrowly to long, thin spermatophore bursa (spb). Sperm gutter continuing Figure 58. Reproductive anatomy of Juga (Oreobasis) new- berryi (USNM_ 1100660). Exte smal, left lateral view of pallial oviduct. Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove, “<4” indicates transition be- tween albumen and capsule glands. Abbreviations: ag, albu- men gland; eg, capsule gland; gg, gonoductal groove; evi, renal oviduct; res, seminal receptacle; sg, sperm gutter; spb, sper- matophore bursa. Scale bar = 1 mm. E. E. Strong and T. J. Frest, 2007 Page 59 posteriorly as shallow groove to small, narrow seminal receptacle (res). Moderate glandular protuberance from opposing inner surface of lateral lamina extending into receptacle aperture. Glandular ridge extending from protuberance to oviduct anterior tip along inner edge of lateral lamina opposite seminal groove; ridge thinning anteriorly. Remarks: The proximal albumen gland forms a promi- nent u-shaped tube that is larger than that in fuga sili- cula. The spermatophore bursa is smaller than both pre- ceding species, but as in J. acutifilosa, the glandular ridge directly opposes the seminal groove and the glandular protuberance is intermediate in size between that of J: silicula and ]. acutifilosa. Apart from minor individual variation in shape and development of the prostate glands, there are no detect- able discrete differences in male reproductive anatomy compared to Juga silicula. Some minor differences in- clude the fact that the ventral fold of the medial lamina appears less textured as in J. acutifilosa, and the dorsal posterior flange appears consistently larger. DISCUSSION EVALUATION OF MORPHOLOGICAL CHARACTERS: Most anatomical differences among the three species investi- gated herein comprise qualitative variations in size and/ or shape (e.g. operculum, hypobranchial gland, oviposi- tor, radula, and bladder). Although often emphasized in species-level systematic studies of gastropods, male and female reproductive anatomy also displays only minor differences in shape and size of the various glands, pouches, and gutters. Additional sampling within and be- tween Juga species is necessary to determine if any of these comprise discrete rather than continuous character variation. Surprisingly, the midgut offers a significant source of variation, with medincatons to size of the cae- cum, style sac, and proximal intestine, as well as overall proportions of the gastric chamber and sorting area. This is very different from eastern North American pleuro- cerids with species from disparate genera displaying al- most identical midgut morphologies (Strong, 2005; Strong, unpublished data). For a summary of ‘these and other differences, see Table 1 The only published anatomical account of any North American Juga is that of Prozorova and Raschepkina (2004) on the female reproductive anatomy of five un- determined Juga species from the Willamette River sys- tem in Oregon. That study indicated the size and shape of the seminal receptacle and spermatophore bursa can vary, which was confirmed in the present study. How- ever, Prozorova and Raschepkina reported seve ral addi- tional findings that could not be confirmed here, includ- ing a sperm gutter that becomes suddenly shallow (an- terior one fifth to one sixth), variable le neth of the opening between the gonoductal groove and momen cav- ity, and asymmetrical arrangement of albumen and cap- sule glands. Indeed, the dorso-ventral axis (as defined by a plane extending between the gonoductal groove and the opening to the mantle cavity), is not correctly iden- tified by these authors. Thus, they mistakenly conclude that the lateral lamina is glandular and the medial lamina is non-glandular, comprising the spermatophore bursa and sansivial receptacle. Consequently, the glands of the oviduct are misinterpreted as a solid block penetrated by channels within the lateral lamina, rather than as a con- voluted tube. This misunderstanding does not allow them to correctly identify the basic symmetry between the medial and lateral laminae, and the proportional de- velopment of the capsule and albumen glands along the antero-posterior axis. Instead, the albumen gland is often depicted as extending far anteriorly, dorsally overlying the c capsule g gland. Not only is this i incorrect, it is difficult to imagine how such an arrangement would function. ANATOMIC AND SYSTEMATIC AFFINITIES OF [UGA: — No conchological feature unambiguously distinguish res Juga from eastern North American pleuroce rid genera; in- deed, Juga has often been synonymized w ith lamin (as Goniobasis )—a genus wide spread east of the continental divide (e.g. Tryon, 1865, 1873; Pilsbry, 1899; Walker, 1918; Henderson: 1935a, b; Goodrich, 1942). However, increasing evidence not only supports the independence of Juga, but suggests ties to Asian pleurocerids. In an analysis of a large segment of the mitochondrial 16S rRNA gene including representatives of five eastern North American pleurocerid genera, Juga falls to the base of the tree rooted on Melanopsis praemorsa (L.) (Melanopsidae) and Melanoides tuberculata (Miiller, 1774) (Thiaridae) (Holznagel and Lydeard, 2000). In an analysis of cerithioidean relationships using nearly full length 16S rRNA sequences (Lydeard et al., 2002), East- erm North American pleurocerids ( (Elimia, Pleurocera) are supported as more closely related to Melanopsis than to a clade uniting Juga with Eastern Asian pleurocerids (Semisulcospira Boettger, 1886, Hua Chen, 1943). Available evidence from reproductive anatomy may also support a link between fuga and Asian pleurocerids. A recent contribution by Strong (2005) re-described the anatomy of Pleurocera acuta Rafinesque, 1831 and Elimia livescens (Menke, 1830) with a re-evaluation of pallial oviduct homologies. Despite several erroneous ac- counts (Woodard, 1934; Jones and Branson, 1964; Dazo, 1965), Strong’s (2005) analysis confirmed Eastern North American pleurocerids described thus far lack a seminal receptacle—a feature present in all described western North American ( (Juga) ) and Asian (Hua, Se misulcospira) species (Itagaki, 1960; Nakano and Nishiwaki, 1989; Pro- zorova, 1990; Rashchepkina, 2000; Prozorova and Ra- schepkina, 2001, 2004, 2005). However, as stated above, the only information available on the anatomy of Juga concerns female reproductive anatomy. Given the de- tailed description of three Juga species herein, it is now possible to better place this unique assemblage within the emerging anatomic and phylogenetic framework for limnic cerithioideans. Page 60 THE NAUTILUS, Vol. 121, No. 2 Table 1. Summary of anatomical differences between three species of Juga. Juga (Juga) silicula Juga (Calibasis) acutifilosa Juga (Oreobasis) newberryi External Anatomy: Operculum nucleus size (as percent of total length) Ovipositor pore Hypobranchial gland Alimentary System: Dentition: Rachidian tooth Lateral teeth Marginal teeth Rachidian basal denticle Rachidian basal margin Weakly formed outermost denticle of lateral teeth Radular sac Salivary glands Sorting area Crescentic groove Caecum Style sac Proximal intestine at base of style sac Reno-Pericardial System: Bladder excretory tubules Reproductive System: Proximal albumen gland Spermatophore bursa Sperm gutter Lateral lamina glandular ridge Prostate glands 45% Shallow Transversely ridged 3/1/38 2/1/2-3 6/7 Present Bluntly v-shaped Present Short Thick, tubular Elongately rectangular Shallow, broad Shallow Large Large protuberance Loosely and regularly branched Moderately large, u-shaped tube Large, rounded Anterior one fourth of oviduct: extends to seminal receptacle as shallow groove within medial lamina Inside seminal groove; large protuberance extending into receptacle aperture Tightly interlocking 41% Deep, highly ridged Thin 2/1/2, 2/1/34 5/6 Lacking, slightly developed Concave bordering rounded median projection Absent Long Thin, tubular Elongately triangular Shallow, narrow Deep Large Small protuberance Loosely and regularly branched Small, rounded pouch Moderately large, rounded Anterior one third of oviduct; becomes obsolete before reaching receptacle Opposite seminal groove; low, flat protuberance extending into receptacle aperture Loosely interlocking 4 34% Shallow Warty 2/1/2 34/1/34 4-5/6-7 Lacking Bluntly v-shaped Present Moderately long Thick, tubular Broadly triangular Shallow, broad Shallow Small Large protuberance Densely and highly branched Large, u-shaped tube Narrow, tubular Anterior one third of oviduct: extends to receptacle as shallow groove within medial lamina Opposite seminal groove; moderate protuber rance extending into receptacle aperture Tightly interlocking Juga, like other pleurocerids, possesses an ovipositor involved in the deposition of the egg capsules (except the viviparous Semisulcospira; Itagaki, 1960) (e.g. Jewell, 1931; Woodard, 1934; Magruder, 1935b; Morrison, 1954; Jones and Branson, 1964; Prozorova, 1990; Rash- chepkina, 2000; Prozorova and Raschepkina, 2001, 2004 2005; Strong, 2005). The ovipositor pore in Juga forme a broad, shallow triangular shelf with a highly grooved epi- thelium. This is very different from the deep pore that expands medially into the foot of Plewrocera and Elimia, with parallel folds that direct the movement of ova through the pore (Strong, 2005). Indeed, this feature was cited by ” ae (1966) as justifying the independence of Juga.U nlike Juga, the distal ovipositor groove extends to the foot sole in some but not all of eastern North Ameri- can pleurocerids (e.g. Van Cleave, 1932; Morrison, 1954; Strong, 2005): ovipositor morphology of Asian pleurocer- ids is unknown. In melanopsids, the pore is deep, glan- dular and complex and the distal 1 groove does not ye sect the foot sole (Bilgin, 197: 3. Glaubrecht, 1996) (Ta- ble 2) Similar to other pleurocerids, the gut of Juga species is characterized by the presence of tubular salivary glands that pass through the nerve ring, the absence of a mid- esophageal eland, and a style sac in restricted communi- cation with the proximal intestine (Mz agruder, 1935a, b: Itagaki, 1960; Dazo, 1965; Strong, 2005). Rachidian basal eae *s are present in at least some Asian pleurocerids (Ko et al., 2001) and are apparently lacking in many eastern North American ple surocerids (e. g. Minton et al., 2004; Sides, 2005). However, they are easily overlooked in whole mounts and their absence from existing descrip- tions may be an error; for example, basal denticles are present in Elimia livescens (Glaubrecht, unpublished E. E. Strong and T. J. Frest, 2007 Page 61 Table 2. Summary of anatomical differences between Juga and other limnic gastropods classified in the Pleuroceridae and Mel- anopsidae. Details from Sunderbrink, 1929; Sods, 1936; StarmiihIner and Edlauer, 1957; Itagaki, 1960; StarmiihIner, 1970; Bilgin, 1973; Houbrick, 1958; Nakano and Nishiwaki, 1989; Glaubrecht, 1996; Strong and Glaubrecht, unpubl. data. NA = not applicable. Elimia livescens Juga Pleurocera acuta External Anatomy: Ovipositor pore Alimentary System: Salivary glands Salivary gland position Esophageal gland Digestive gland ducts Caecum Reno-Pericardial System: Bladder Evagination of bladder wall Shallow, simple, weakly glandular Tubular Dace ‘ Pass through nerve ring Absent ] Shallow/Deep Small, pallial Present Deep, simple, Semisulcospira Melanopsidae NA Complex, highly weakly glandular glandular Tubular Tubular/branched Tubular Pass through nerve ring Pass through nerve ring Pass through/anterior to nerve ring Absent Absent? Present 2 2 | Shallow Shallow Deep and spiral Small, pallial ? Small, pallial Absent ? ? Nervous System: Dialyneury, Zygoneury Zygoneury Zygoneury Dialyneury? Zygoneury Accessory ganglion Present Present Absent? Present? between left pleural and sub-esophageal ganglia Reproductiv e System: Ovipositor distal groove Present Present/Absent NA : Absent intersects foot sole Seminal vesicle Straight Straight Straight Folded Proximal albumen gland U-shaped Straight 2 ? Gonoductal groove Convoluted Simple Simple? Simple? Seminal receptacle Present Absent Present Present Reproductive strategy Oviparous Oviparous Viviparous Oviparous data), but have not been described in the literature (e.g. Baker, 1928; Dazo, 1965). The phylogenetic significance of these features is unclear as they occur sporadically among many cerithioidean groups, including batillariids, planaxids, melanopsids, thiarids (see e.g. Thiele: 1928; Houbrick, 1987: Glaubrecht, 1996). Melanopsids differ in eee an esophageal gland, and the salivary glands may be tubular or branched and variably pass through or by-pass the nerve ring; the radula is variable and may present a rachidian ee is similar to that of pleuroce rids, or may be quadrangular, robust with conical denticles and with a marked glabella (Sunderbrink, 1929; Thiele, 1928: Bilgin, 1973; Glaubrecht, 1996). The midgut of Juga species diverges from eastern North American pleurocerids and other putatively closely related limnic lineages most noticeably in size and shape of the glandular pad ‘and configuration of the cres- centic ridge. In Juga, the glandular pad is rather narrow with a crescentic ridge hata is often separated from it by a wide, shallow groove; the proximal end of the crescen- tic ridge borders a vestibule that receives several ducts of the digestive gland. Typically, melanopsids, paludomids, and thiarids have a broadly rounded glandular pad and a deep, narrow crescentic groove such that the crescentic ridge closely adheres to the outer edges of the pad. Al- though the midgut has demonstrated great utility in re- constructing relationships among seatioiices line ages (Strong, unpublishe d data), midgut characters of Jiga do not provide unambiguous evidence of affinity to any one freshwater family; the small, narrow glandular pad is rather similar to that of Se misulcospira, "bist the configu- ration of the crescentic ridge, particularly in J. silic ae and J. newberryi, is unique among cerithioideans known thus far. The presence of a single digestive gland duct vestibule is shared between Juga and melanopsids, but is also found in a number of cerithioideans; other pleuro- cerids and paludomids have two digestive gland ducts (Strong and Glaubrecht, 2002, 2003, 2007, unpublis! ied data; Strong, 2005). As in paludomids ( e.g. Strong and Glaubrecht, 2002, 2003) and melanopsids ( (Bilgin, 1973), the kidney of pleu- rocerids penetrates the palli: ul cavity (Mz agruder, L935b; oe 1960: Strong, 2005), but the pallial portion of the bladder is smaller Than that in the former taxa. The branching pattern of excretory tubules within the blad- der is essentially identical between Juga and eastern North American pleurocerids, but Juga is unique in the outpocketing of the wall separating the main chamber and the bladder (Strong, 9005) Kidney anatomy is cur- rently unknown for Asian pleurocerids ‘and me lanopsids Page 62 THE NAUTILUS, Vol. 121, No. 2 Although published accounts of pleurocerid nervous systems disagree on the number of nerves produced by various ganglia (Magruder, 1935b; Itagaki, 1960; Dazo, 1965; Strong, 2005; present study), this is often variable within species. However, the present account agrees with that of Strong (2005) that the cerebral ganglia pro- duce seven nerves, and that there are two prominent anterior pedal nerves with a variable number of small accessory nerves (typically four to seven). The thickened connective between the left pleural and sub-esophageal ganglia, representing a small accessory ganglion, was found to give off only a single large nerve in the present study, bar was found by Strong (2005) to produce one to three nerves in Elimia and Pleurocera. Str ong (2005) also found the sub-esophageal ganglion to give off one to three nerves, whereas three to four nerves were found in the present study. The most intriguing aspect of the nervous system is the presence of the accessory ganglion. It is known only from eastern North American Sead (Magruder, 1935b: Strong, 2005) and now Juga (present study) ), and has not been depicted in most accounts of melanopsid (Bouvier, 1887: Sods, 1936; Starmiithlner and Edlauer, 1957; StarmiihIner, 1970; Bilgin, 1973; Glaubrecht, 1996) and Asian pleurocerid Gnesi, 1960) nervous system anatomy. However, the illustration of the nerve ring of Melanopsis frustulum Morelet, 1856-57 (St: ee 1970) and that of M. doriae Issel, 1866 (StarmiihIner andl Edlauer, 1957) clearly show a thickened connective be- tween the left pleural and sub-esophageal ganglia, indi- cating that it is most likely present in melanopsids as well. Paludomids and thiarids have a much more con- centrated nerve ring with the left pleural and sub- esophageal ganglia fused or in close contact (e.g. Se- shaiya, 1934: Glaubrecht, 1996; Strong and G laubrech® 2002. 2003). Pleurocerids and melanopsids share the same basic layout of the pallial oviduct to the exclusion of paludo- mids and thiarids, including the presence of a long open- ing to the mantle cavity, anda deep sperm gutter open- ing anteriorly to a spermatophore bursa and posteriorly to a seminal receptacle; as mentioned above, eastern North American pleurocerids are unique in lacking the seminal receptacle (Bilgin, 1973; Nakano and Nishiwaki, 1989: Prozorova, 1990: Glaubrecht, 1996: Rashchepkina, 2000: Prozorova and Raschepkina, 2001, 2004, 2005: Strong, 2005). Whereas in Juga and Semisulcospira the sperm gutter becomes obsolete or continues posteriorly as a shallow groove within the medial lamina (Prozorova and Aaschepkina, 2004, 2005; herein), in melanopsids the sperm gutter is continuous along the ventral edge of the medial lamina and contains the opening to the re- ceptacle at the posterior end of the oviduct (Bilgin, 1973; Glaubrecht, 1996). In Elimia and Plewrocera, a deep sperm gutter 1s present above the opening to the mantle cavity along its entire length, and closes posteriorly to form a short, blind spermatophore bursa. One aspect of cerithioidean reproductive anatomy that is routinely overlooked is the configuration of the capsule and albumen glands. Strong and Claus echt (2002, 2003) have found that the shape of the albumen gland in palu- domids is quite distinctive, and likely a synapomorphy of the family. In fuga, the glands and intervening gonoduc- tal groove of the pallial oviduct are highly conimlued with a proximal albumen gland that is pouch- -like or u- shaped (present study), elas eastern North American pleurocerids possess ¢ clands that form two narrow bands with smooth opposing surfaces and an essentially linear proximal albumen gland (Strong, 2005). However, such potentially informative characters are undescribed for other pleurocerids and melanopsids. Male reproductive anatomy of Juga is apparently quite distinct as no other male pleurocerid (indeed, no other cerithioidean) has been described with a tightly inter- locking arrangement of glands (e.g. Woodar d, 1934; Ita- gaki, 1960; Prozorova, 1990): however, a large fold within the lateral lamina may be present (Nakano and Nishi- waki, 1989; Strong, 2005). Among eastern North Ameri- can pleurocerids, hoch the highly Folded proximal portion (Woodard, 1934) and distal region of the prostate Strong (2005) have been inferred as the site of spermatophore formation. In Juga, there is no differentiated anterior or posterior region, but intuitively the tightly interlocking folds of the Tateral and medial laminae must function in molding the spermatophore. CONCLUSIONS While sharing many similarities, numerous features clearly set Juga apart from eastern North American pleu- rocerids: ovipositor pore, lateral outer cusps, midgut glandular pad and crescentic ridge, digestive gland duct vesubule; evagination of kidney "wall: convoluted gono- ductal groove, proximal albumen gland, seminal recep- tacle, interlocking prostate glands. However, as is often the case for ancient lineages , it is difficult to identify uniquely shared features that more or less unambigu- ously demonstrate affinity to any one limnic lineage. Juga is particularly difficult as many of these features not only set the genus apart from other pleurocerids, but are un- dccumente d (ovipositor pore, kidney, pallial oviduct glands) or apparently autapomorphic among cerithioide- ans (crescentic ridge, evagination of the kidney wall, prostate). Nevertheless, this analysis has revealed some characters that are shared with Asian pleurocerids (mid- gut glandular pad, seminal receptacle), but some that are also shared with melanopsids (digestive gland duct). However, the presence of the seminal receptacle is un- doubtedly plesiomorphic and uninformative in delineat- ing rele tionships. Others are so heterogeneously distrib- uted among limnic lineages, it is difficult to determine if there may be an unde rlying phylogenetic signal (oviposi- tor ventral eroove, Ti achidian basal denticles), Unlike mo- lecular dats. given the available morphological evidence, there is no overwhelming signal linking Juga to Asian pleurocerids, nor for that matter linking eastern North E. E. Strong and T. J. Frest, 2007 Page 63 American pleurocerids to melanopsids (Table 2). While part of this pattern may be due to the high rate of auta- pomorphies, it may simply be an artifact of missing data. Clearly, one of f the more significant impediments in as- sessing pleurocerid affinities is that comprehensive ana- tomical accounts of pleurocerids and melanopsids are rare, leaving many potentially informative characters un- known. Thus, it is clear that further anatomical studies within the context of ongoing morphological and molecu- lar cladistic analyses are necessary to unambiguously re- solve the question of pleurocerid monophyly and their affinities to melanopsids. These results also have implications for the systematics of Juga, particularly with regards to the y validity of cur- rent subgeneric subdivisions. Although all three taxa pos- sess unique features that clearly distinguish them from one another, ]. silicula and J. newberryi are strikingly anatomically similar and share many features to the ex- clusion of J. acutifilosa: 1) shallow, weakly ridged ovi- positor pore, 2) similar rachidian basal margin, 3) similar development of weak, outermost denticle of the lateral teeth, 4) thick, tubular salivary glands, 5) shallow, broad crescentic groove, 6) shallow caecum, 7) large proximal intestine protuber rance on base of style sac, 8) u-shaped proximal albumen gland, and, 9) tightly interlocking prostate glands (T Table 1). In conclusion, the enatomieal data do not support separation of Juga sensu stricto and Oreobasis and Oreobasis is thus here synonymized with Juga sensu stricto; Calibasis is sufficiently distinct to merit retention as a valid taxon. The question of the rank of these taxa is, of course, highly subjective. But given what is known about Anotouical differentiation among other pleurocerid genera, it could reasonably be ar cued that Juga sensu stricto and Calibasis be recognized at the rank of genus. ACKNOWLEDGMENTS We are indebted to Marilyn Schotte (USNM) for inking the anatomical drawings and for scanning electron mi- oa We also thank Philippe Bouchet (Muséum na- tional d'Histoire naturelle, Paris) for critical reading of the manuscript and consultation on matters of zoological nomenclature. Arthur Bogan (North Carolina State Mu- seum of Natural Sciences) and an anonymous reviewer provided valuable comments. We are also grateful to Ed Johannes (Deixis Consultants) for Figure 1, for collecting specimens with TF, and for compiling locality descrip- tions. Support to Deixis Consultants from the Cantara Trustee Council is gratefully acknowledged (Grants C0010010, CO110013). LITERATURE CITED Baker, F. C. 1928. The fresh water Mollusca of Wisconsin. Part I. Gastropoda. 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Journal of the Tennessee Acad- emy of Science 9: 243-259, THE NAUTILUS 121(2):66-75, 2007 Page 66 Discriminating shells of Gastrocopta pentodon (Say, 182 2) and G. tappaniana (C. B. Adams, 1842) (Gastropoda: Pulmonata) with an example from the Delmarva Peninsula, eastern USA Timothy A. Pearce! Marvin C. Fields Carnegie Museum of Natural History 4400 Forbes Avenue Kayoko Kurita National Institution for Academic Degrees and University Evaluation 1-29-1 Gakuen-nishimachi, Kodaira-shi Pittsburgh, PA 15213 USA Tokyo, 187-8587 JAPAN ABSTRACT The North American pupilloid land snails Gastrocopta pent- odon and G, tappaniana have similar shells that can be difficult to separate, which raises the question of whether they repre- sent two species or environmentally influenced variants of one species. In 1906, Vanatta and Pilsbry presented 53 drawings of shells of the two species to facilitate separation, but differences therein were subtle. Discriminant function analysis of measure- ments from their illustrations gave a discriminant function clas- sifying 96% correctly, with aiaimnal overlap between groups shown by factor analysis. The function revealed both forms on the Delmarva Peninsula, again with minimal factor analysis overlap. Bimodality of morphology does not reflect sexual di- morphism in these hermaphrodites. The two forms maintained their distinct morphologies where they co-existed, supporting the concept of separate species. Reports of the forms in habitats of different wetness could indicate separate species with different moisture preferences or one species with moisture-influenced morphology. We found the two forms to show distinct morphologies in medium wet areas, further supporting the concept of separate species. Regarding habitat wetness, we confirmed Delmarva Gastro- copta tappaniana in wetter areas, whereas G, pentodon oc- curred in a wide range of moistures, but tending to be found in drier areas. Our surrogate measure of habitat wetness relying on plant moisture requirements should be useful in future stud- ies. Geographically, G. tappaniana tended to occur along the SE Delmarva Atlantic coast while G. pentodon ranged more wide ly In an application of the discriminant function, measurements from an image of the lectotype of Gastrocopta carnegici clas- sified that species with G. pentodon. INTRODUCTION The ability to distinguish one species from another is central in biology. However, environmental variation in Corre sponding author pearcet@carnegiemnh org form can sometimes be mistaken for species-level differ- ences in taxa including mollusks (Minton and Gunder- son, 2001). Historically, making a distinction between the North American pupilloid land snails Gastrocopta pentodon (Say, 1822) and Gastrocopta tappaniana (C. B. Adams, 1842) has been difficult (Vanatta and Pilsbry, 1906). These latter authors discussed and evaluated G. pen- todon, G. tappaniana, and other similar, previously de- scribed forms. They also illustrated various modifications of form, Their paper presented drawings of 53 Gastro- copta specimens from eastern North America. Although Vanatta and Pilsbry identified some of their illustrated specimens as G. pentodon and others as G. tappaniana, they did not reveal their criteria for specific allocation of these individual specimens. Vanatta and Pilsbry (1906) and Pilsbry (1948) de- scribed distinctions between Gastrocopta pentodon and G. tappaniana including differences in shell size and shape, and the number and arrangement of apertural teeth. Despite Vanatta and Pilsbry’s clarification of the species differences, confusion persists and some workers have consolidated the two forms into one single species while others have kept them as separate species. Bequaert and Miller (1973: SS-90) lumped the species, stating that the holotype (actually the lectotype, selected by Clench, 1965) of G. tappaniana is a typical G. pen- todon. On the other hand, Hubricht (1976: 107) also examined the type of G. tappaniana and concluded it was G. tappaniana and not G. pentodon. Lauriol et al. (2003) and Nekola (2004) recognized the two forms as separate species. Our visual e xamination of Vanatta and Pilsbry’s (1906) drawings of the 53 specimens in light of their statements comparing the os species left us uncertain that those shell characters could reliably separate the two species. Some of the G. pentodon seemed to have characteristics of G. de tarlge G vice versa (see in particular their figs. 17, 32, 45, . Admittedly, Vanatta and Pilsbry at- tempted ie aha variability in the forms, such that they might have chosen extreme examples. T. A. Pearce, M. C. Fields, kK. Kurita, 2007 Page 67 Multivariate statistical procedures are obviously more accessible now than they were in 1906. Factor analysis (FA) identifies the axes of mé yor variation in a dataset and helps show whether specimens cluster into more than one group in a multidimensional space. Discrimi- nant function analysis (DFA) can identify which variables are most useful for separating two pre-defined groups and can identify coefficients to use with the variables in a discriminant function (DF) for classifying unknown specimens. Nekola and Barthel (2002) use da r simlae ap- proach. Some researchers have suggested that Gastrocopta pentodon prefers drier areas and G. tappaniana prefers moister areas (Sterki, 1906: 134; Pilsbry, 1948: SSS—S90; Hubricht, 1985: 9: Nekola, 2004). One hypothesis holds that the forms are separate species as evidenced by their different moisture preferences. An alternate hypothesis is that, since larger snails are sometimes associated with moister conditions (Goodfriend, 1986, and references therein), the two forms might simply be two ends of an environmentally influenced continuum of morphology. If environmental conditions are continuous, then randomly sampled specimens should show continuous morphological variation if they represent one environmentally influenced species or a bimodal distribution if they are two species with different moisture preferences. F urthermore, examin- ing w hether the two species maintain their separate mor- phologies where they co-occur would provide strong evi- dence whether they are two separate species. Numerous specimens tentatively identified as Gastro- copta pentodon were available from a survey of terrestrial mollusks on the Delmarva Peninsula (Pearce and Italia, 2002). Examining these specimens using morphometric methods should address whether both forms occur on Delmarva and, if so, whether the forms are distinct and whether they differ in their moisture preferences. In this paper we explore the moisture associations of the two forms on the Delmarva peninsula to examine whether specimens identified as G. pentodon occurred in drier habitats and whether G. tappaniana occurred in moister habitats. Furthermore, we look for interpretable differences in geographical distribution of the two forms on the peninsula. Finally, by examining specimens at localities where the two forms co-occur, we address whether the two forms are valid species or ecomorphs of a single species. We address three principal questions in this paper: (1) Are the two forms identified by Vanatta and Pilsbry (1906) distinguishable via morphometric analysis and, if so, which fenpines discriminate between them? To an- swer this question, we examined selected variables from their 53 illustrations in FA and DFA. Do the Gastrocopta pentodon and G. tappaniana from the Delmarva Peninsula fall into two morphometric groups and, if so, how do the features of those two groups compare with those of the forms as identified by Vanatta and Pilsbry (1906)? To answer this question, we applied the DF from question (1) to 577 Delmarva specimens and examined results along FA axes. (3) Does the wetness of the habitats of the Gastrocopta pentodon and G. tappaniana forms differ as previously suggested? How are they distributed on the peninsula? Does co-occurrence evidence addresses whether the two forms represent separate species or environmentally in- fluenced morphs of the same species? To answer this question, we compared occurrences using wetness esti- mated from plant data and compared morphology of the forms in sympatry and allopatry. MATERIALS AND METHODS Sources of Specimens: We examined illustrations of the 53 Gastrocopta shells presented by Vanatta and Pils- bry (1906), which had been drawn to the same scale oe using a camera lucida. Their specimens were from geographically widespread areas in eastern North America. They identified the specimens as 41 G. pen- todon and 12 G. tappaniana. We examined 577 unbroken adult Gastrocopta shells from 130 leaf-litter samples from 123 grid squares on the Delmarva Peninsula. The number of shells measured per sample ranged from 1 to 69. The Delmarva Peninsula shells came from a study of land snails on the peninsula (Pearce and Italia, 2002) in which leaf litter samples (1-4 liters) were collected from 794.5 « 5 km grid squares, passed through sieves, and snails retained by screens 0.5 mum or larger mesh were picked and identified. We ex- cluded juveniles (adults are easily recognized by the presence of a reflected lip and we I-deve loped apertural teeth) to avoid introducing variability from shells of dif ferent ages and excluded fe other clearly different spe- cies of Gastrocopta found on Delmarva. Although speci- mens resembling G. pentodon and G. tappaniana were found in 201 samples from 11 different grid squares on Delmarva (Figure 1) excluding broken, juvenile, or wet- preserved specimens (bodies of alcohol-preserved speci- mens obscured apertural features) reduced the number of measurable specimens. Differences between the datasets: the Vanatta and Pilsbry geographic coverage is about two orders of mag- nitude larger (throughout « -astern North America for Vanatta and Pilsbry, just the Delmarva Peninsula for Delmarva), whereas the Delmarva sample size is an or- der of magnitude larger (n = 53 for Vanatta and Pilsbry, n = 577 for Delmarva). Voucher specimens are deposited at the Delaware Museum of Natural History. Selection and Measurement of Variables: \Ve took measurements from camera lucida drawings of shells, using 53 drawings by Vanatta and Pilsbry (1906) and 577 of our own drawings of specimens from the Delmarva Peninsula. Page 68 THE NAUTILUS, Vol. 121, No. 2 Figure 1. Delmarva Peninsula showing locations where specimens of Gastrocopta pentodon and G tappaniana were found We measured or counted 8 variables from the draw- ings and derived 13 additional (ratio) variables as com- binations of the measured variables (Table 1, Figure 2). We excluded teeth on the parietal wall from the count of teeth because whereas only one parietal tooth is present in Gastrocopta tappaniana, either one or two parietal teeth can be present in G. pentodon (Pilsbry, 1948: 889). \perture height was measured from the middle of the parietal callus. We included derived variables because ratios suc- cinctly describe shape, which is not described by original measurements. Because ratios are more intuitive, includ- ing ratios in this analysis should give a more useful result for separating these species. Although ratios can theo- retically produce non-normal distributions, the ranges of our variables are limited so they can be used in FA and DFA. We did not check for normality or transform vari- ables because FA and DFA are robust against non- normality (Mardia, 1971; Hagiuda and Shigemasu, 1996). In order to reduce she fcmabe sr of var bles to about 1/5 the number of specimens (Hair et al., 1998), and to determine the set of variables that have the greatest chance of separating the species as defined, we used non-parametric Mann- Whitney U-tests to compare mea- surements of each variable on the two Gastroc ‘opta forms as defined (Table 1) and used the variables that showed significant differences between the two groups in further analyses. Statistical Techniques: DFA and FA were per- formed using SAS version 8.02 (SAS Institute Inc., Cary, NC). We used DFA to determine the variables most useful in discriminating between the two forms using stepwise, backward, and forward methods for variable selection. The three methods gave similar sets of vari- ables to use in separating the species, giving us confi- dence that two recognizable groups exist in the data set. Then SAS applied the DF to the analyzed specimens to determine reclassifications and percent correct classifi- cation. We applied those DF coefficients to measure- ments of the 577 specimens from the Delmarva Penin- sula to determine which forms occur there. We used FA to examine graphically whether two mor- phological groups of individuals are evident among the specimens, using the variables determined by the DFA. We used FA with principal compone nts (esse tially the same as principal component analysis with rote ition). We accepted the first three axes, applie sd varimax rotation, and plotted specimens on the factor axes Distinctness of the Forms: To further study distinct- ness of the two forms, in addition to examining FA plots, we examined whether individual samples from Delmarva contained mostly one form of Gastrocopta or if forms tended to be randomly distributed among samples. To examine whether this non-randomness represents two species, or one environmentally influenced species, we examined their morphologies in symp yatry. If the forms are two morphologically distinct species, then their mor- phologies should remain distinct and mixed samples, containing both forms, should exhibit bimodal morphol- ogy. If the forms are a single species with environmen- tally de pendent morphology, morphology should not show bimodality under intermedi: i: environmental con- ditions; in intermediate conditions, morphology should be intermediate. To verify the names being applied to the two forms, we measured and compared the lectotype of Gastrocopta tappaniana, To examine visually how it compares with the other specimens, we plotted the pseudo-factor scores (calculated from lectotype measurements using results of FA on the 53 specimens, rather than including it in an analysis with the other 53 specimens) for the lectotype where it would appear on the factor plots gene rated for T. A. Pearce, M. C. Fields, K. Kurita, 2007 Page 69 Table 1. Variables recorded on Gastrocopta pentodon and G. tappaniana. Derived variables relate to shape. Bold p-values indicate variables showing significant difference (Mann-Whitney U test) between forms as identified by Vanatta and Pilsbry (1906). Measured variables are in millimeters. Variable Type Description p-Value between forms aph/bdh Derived Aperture height/body whorl height 0.6863 aph/h Derived Aperture height/shell height 0.0629 aph Measured Aperture height 0.0003 bdh Measured Body whorl he ight 0.0001 bdh/h Derived Body whorl ht/shell ht 0.0145 bdhAwv Derived Body whorl height/shell width 0.1948 h Measured Shell ht 0.0660 hat Measured Hat height (height above penultimate whorl) 0.2464 hat/h Derived Hat height/shell height 0.0029 hat/spr Derived Hat height/spire (spire = hat height + penultimate 0.0007 whorl height) pnilth Measured Penultimate whorl height 0.0126 pnlth/bdh Derived Penultimate whorl height/body whorl height 0.6100 pnith/h Derived Penultimate whorl height/shell height 0.2177 pnlth/hat Derived Penultimate whorl he ight/hat 0.0007 pulth/pnltw Derived Penultimate whorl he .ight/penultimate whorl width 0.9915 pnitw Measured Penultimate whorl width 0.0002 pnitw/h Derived Penultimate whorl width/shell height 0.0742 pnitw/w Derived Penultimate whorl width/shell width 0.7823 tthxpar Counted Number of teeth excluding those on the parietal 0.0000 (upper) wall w Measured Shell width 0.0002 wh Derived Shell width/shell height 0.1313 just the 53 specimens. Second, we performed DFA to determine how it classified. Moisture Association of the Forms: To examine the relationship of the forms with the moisture in their en- vironment, we calculated a surrogate index of wetness using plants noted at the collecting sites. Because plant species differ in their long-term moisture requirements, estimating éaveninental moisture from plant data should provide a biologically meaningful measure of long-term moisture av: ailability. Wetland facultative indicator status of many North American plant species is available at USDA-NRCS (2004) by geographical regions. The plants are assigned to one of five categories depending on how obligate od they are to wet lands. For each plant species at a locality, we assigned a score of 5 for the wettest facultative indi- cator rane 1 to the driest, and 2, 3, or 4 to the three intermediate ranks on the website (adding 0.3 for a “+” and subtracting 0.3 for a “—"). We calculated the mean facultative indicator rank of all the plants recorded at each locality and used this mean as a surrogate index of wetness at the locality. To test the hypothesis that Gastrocopta pentodon oc- curs in drier and G. tappaniana in moister areas we di- vided the wetness score for specimens into three parts, with the middle part representing 25% of the specimens, and the other two parts being approximately equally di- vided. Then we examined the number of individuals in each of the three wetness areas as a function of their DF scores. Geographical Distribution of the Forms: We ex- amined a map of localities of specimens identified by the DF to see whether the two forms tended to show inter- pretable geographic patterns on the peninsula. Of the 130 samples, 106 were only Gastrocopta pentodon or G. tappaniana; the 16 that were predominantly G. pentodon were scored as G. pentodon, the 2 that were predomi- nantly G. tappaniana were scored as G. tappaniana, and we omitted the 5 samples containing equal numbers of the two forms. We recognize that geographical location and wetness might not be independent. RESULTS Distinctness of 53 Specimens: DFA of the 53 speci- mens as defined by Vanatta and Pilsbry (1906) deter- mined coefficients for four ee to separate the two forms of Gastrocopta (Table 2). To use these coefficients to determine an unknown spe cimen, first multiply each DF coefficient by the corresponding variable value for that specimen, and then sum all these results. If the result is negative, then the specimen is more likely G pentodon, and if greater than zero, the specimen is more likely G. tappaniana. For example, specimen 1 in Vanatta and Pilsbry (1906) has the following measurements (mm)/count: bdh = 0.942. hat/h = 0.267, pnith/hat = 0.645, tthxpar = 4. Multiplying these ae by the DF coveeut sad adding them yields —8.068, indicating that the specimen classifies as G. pentodon. The DF correctly classified 51 (96.2%) of the 53 speci- Page 70 THE NAUTILUS, Vol. 121, No. 2 tthxpar Figure 2. aperture height [measured from base of outer lip to midpoint of callus connecting the two lip insertions], bdh = body whorl Variables measured on Gastrocopta spp.: aph = height, h = shell height, hat = height of shell above penulti- mate whorl, pnlth = penultimate whorl height, pnltw = pen- ultimate whorl width, tthxpar = number of apertural teeth excluding those on the parietal (upper) wall, w = shell width. Dimensions are measured perpendicular or parallel to axis of coiling. mens that had been used to create the function: 39 of the 41 Vanatta and Pilsbry-identified G. pentodon classified as identified, and all 12 of the 12 G. tappaniana classified as identified. In FA, of the 53 specimens, the first three eigenvalues using the four variables explained 98.1% of the variation. Two shape variables (hat/h, pnlth/hat) loaded strongly on factor 1, one size variable (bdh) loaded strongly on factor 2, and o one variable (tthxpar) loaded strongly on factor 3 (Table 3). This result, in which shape loaded strongly on factor . contrasts with the usual FA pattern in ich factor | is size and factor 2 is shape. The four variables were all important for explaining variation in the dataset, as evidenced by high communality scores (sum of Table 2. Discriminant function (DF) coefficients for separat- ing the two forms of Gastrocopta using four variables Variable Coefficients Constant =fi071] bdh 26.82807 hat/h 70 pnilth/hat 33.85039 tthxpat 0.92107 Table 3. Loadings for the four variables on the three factors from factor analysis (FA) of 41 Gastrocopta pentodon and 12 G. tappaniana specimens illustrated by Vanatta and Pilsbry (1906), using the rotated factor method. Bold loadings indicate the main variables loading on each factor. Percent variance explained is after varimax rotation. Factor | Factor 2 Factor 3 pnilth/hat 0.96674 0.00425 0.16676 hat/h -0.96086 —0.08634 —0.17608 bdh 0.0465] 0.98561 0.16166 tthxpar 0.21990 0.17578 0.95955 Variance explained 1.908 1.010 1.006 squares of factor loadings [SS], ranging from 0.961 to 1.000). Figure 3 plots specimens on factors 1 and 2, and factors 2 and 3, showing that G. tappaniana (solid squares) occupies a portion of morphospace at the edge of and somewhat overlapping with G. pentodon (hollow diamonds). Thus, although the two forms overlap, they generally occupy separate portions of morphospace. Shells scoring higher on factor 1 have a relatively smaller hat to otal height, and a relatively taller penul- timate whorl relative to hat height. Shells scoring higher on factor 2 have a larger body whorl height. The third factor reflects aueaber of teeth, and shells classified as Gastrocopta tappaniana had 7 non-parietal teeth, in con- trast to G. pentodon, which had a variable number of teeth. Distinetness of 577 Specimens: — Applying the DF coefficients to the Delmarva specimens classified 475 specimens as Gastrocopta pentodon and 102 as G. tap- paniana. In FA of the 577 Delmarva specimens, the first three eigenvalues using the four variables selected by the DFA explained 98.0% of the variation. Two sh ape variables (hat/h, pnlth/hat) loaded strongly on factor 1, one size variable (bdh) loaded strongly on factor 2, and one vari- able ( (tthxpi ir) loaded strongly on factor 3 (Table 4). The four variables were all important for ¢ epbinne ieee in the dataset, as evidenced by high communality scores (SS ranging from 0.957 to 1. 000). Figure 4 plots speci- mens on factors | and 2, and factors 2 and 3, showing that the two species occupy generally separate portions of morphospace with minimal ove srlap. Interpretation of factors 1 and 2 are the same as for the Vanatta and Pilsbry results. The third factor, largely reflecting number of teeth, shows variability in both forms, but shells classified as Gastrocopta tappaniana tend not to have the minimum number of teeth. The specimens plot in distinct columns in this analysis be- cause number of teeth was discreet: the columns appear more distinct in this analysis than in the analysis of Vanatta and Pilsbry data because teeth loaded much more strongly on factor 3 in this Delmarva analysis. Distinctness in Sympatry and Type Material: The species composition of samples from Delmarva (contain- T. A. Pearce, M. C. Fields, kK. Kurita, 2007 Page 7] 37 : 3 * * a 2 2 | | ° ° | ° a ° ° a -_ 9 -_— g ~ 2" a - oe ean an a an a ° 2 } a bal Py = ° ‘ 5 Pa S 5 0 S90 ° . Mss 5 f © 990 O% : © o° © oo) © ° 7 ge uw 2 6 ° WL } o 8 “1 oo O° ° ° © ; ° | ° : ° 3° % }° - ° 22 , 2 ‘ a ae 37( -3,-+— = 3. SE) — —_ -3 -2 -1 0 1 2 3 -3 -2 -1 0 1 2 3 Factor 1 (shape) Factor 3 (teeth) Figure 3. | Factor analysis (FA) plots of 53 specimens on factors | and 2 (left) and factors 2 and 3 (right). Hollow diamonds represent Gastrocopta pentodon and solid squares are G, pelnaaeie as identified by Vanatta and Pilsbry (1906). The shell image shows, to the right on factor 1, a relatively smaller hat to total height and a relatively taller penultimate whorl relative to hat height: higher on factor 2 shows a larger body w horl he ight; to the oe on factor 3 shows shells with more apertural teeth. Asterisk (* ) indicates position of lectotype of G tappaniana and plus sy mbol (+) indicates position of the lectotype of G. carnegici. Numbers beside shell images correspond to specimen numbers from eae and Pilsbry (1906). ing 5 or more specimens) was non-random. Samples tended to be ean primarily of one species or the other (Figure 5), instead of mostly mixed samples that would be. expected with random mixing. By the DF, 28 of 39 samples were composed of a single : species. In testing whether the two forms maintain their distinctness in sympatry, we examined the FA plot (factors | and 2) of samples from Delmarva that the DF identified to contain both forms (mixed samples). Figure 6 shows good sepa- ration of the two coexisting forms, supporting the idea that the two forms are separate species. The separation of the two forms in the mixed-only samples is as good as the separation in all samples (compare Figures 4 ‘anal 6). We performed two tests to determine whether one of the two morphologically discrete forms we identified on Delmarva corresponds to type material of Gastrocopta Table 4. Loadings for the four variables on the three factors from factor analysis (FA) of 577 specimens from the Delmarva Peninsula, using the rotated factor method. Bold and under- lined loadings indicate the main variables loading on each factor. ; tappaniana (we could not locate type material of G. pen- todon). First, on the FA plots (Figure 3), the lectotype of G. tappaniana was located in the se he of the ‘graph with specimens classified by Vanatta and Pilsbry as G. tappaniana. Second, the lectotype of G. tappaniana was classified as G. tappaniana by the DF. Moisture Association and Geographical Distribu- tion of Forms on Delmarva: The 22 samples having 5 or more specimens with only Gastroc ‘opta pe mntodon las classified by the DF) were from areas drier (wetness factor X = 2.62, sd = 0.46) than the 6 samples having 5 or mone specimens with only G. tappaniana (wetness X = 3.14, sd = 0.30) (t-test, p = 0.006). Mean wetness factor foe all 471 specimens classified by the DF as G. pentodon was 2.59 (sd = 0.44) and that for the 100 G. tappaniana was 2.95 (sd = 0.44, t-test, p < 0.00005). The histogram in Figure 7 shows the number of individuals by their DF score separately for each of three wetness classes. The vast majority of G. fappaniana (positive DF) occur in medium or wet areas. On the other hand, G. pentodon (negative DF) occur in a wide range of moisture, with a tendency to be in drier areas. In samples from medium wet areas, bimodality of forms is evident, with a hint of Factor | Factor 2 Factor 3 —— a siaebes bimodality in the sample s from dry and wet areas. pnith/hat 0.97099 0.13158 0.03672 A map of localities of specimens identified by DF hat/h -0.94900 -0.23783 -0.01636 shows Gastrocopta pentodon widespread throughout the en 022s! 0.97414 pig peninsula but essentially absent from the southeastern tthxpar ; 0 03081 0.00861 0.99948 coast and barrier islands. In contrast, s: unples containing Variance explained 1.895 1.023 1.001] G. tappaniana were most common oe the Atlantic Page 72 THE NAUTILUS, Vol. 121, No. 2 4 3 -2 -1 0 1 2 3 4 Factor 1 (shape) Figure 4. ° oO Factor 2 (size) oO —_ OGD Mo Rocce om Mm ae Ot. 20S aR, 4 -3 -2 -1 0 1 2 3 4 Factor 3 (teeth) Factor analysis (FA) plots of 577 Gastrocopta specimens from the Delmarva Peninsula plotted on factors 1 and 2 (left) and factors 2 and 3 (right). The same variables were used as in analysis of Vanatta and Pilsbry data. Hollow diamonds represent specimens classified as G. pentodon and solid squares those cle issified as G tappaniana. Coast in southeastern Delmarva and closer to major wa- ter bodies (Figure §). Interestingly, four Atlantic coast samples containing 5 or more specimens of only G. tap- paniana were from four of the wettest areas, as deter- mined by the plant moisture associations. DISCUSSION These results indicate that the two rons of Gastrocopta can be considered separate spe cies. DFA successfully discriminated two groups. The 53 specimens in Vani ta and Pilsbry (1906) could be separated using 4 variables and the DF classified 96% of specimens as defined. The two forms were generally distinct by FA de spite, some morphological overlap. When applie d to the 577 Del- marva specimens, the DF identified both species on Del- marva. FA of the Delmarva specimens showed the two species occupying generally separate portions of mor- phospace with minimal ove le ap. 30 ~ 20 = =| fe) O 10 ° ° ro) [=] fo} fo) (=) ro) co) =) . Y oy SS 8 es oO ro) oO fo) -) fo) [=] f-) fo} ai = AQ o +t wo oO i ire) Q Percent of G. tappaniana Figure 5. Proportion of specimens that were Gastrocopta £ I I } ppaniana in the 39 samples containing 5 or more specimens Even stronger evidence of separate species is that the forms matntined their distinct morphologies where they coexist. Gastrocopta pe ntodon and G. tappantana from the Delmarva Peninsula showed no evidence of conver- gent morphology in sympatry. Moreover, the forms maintained their separate morphologies in samples from 5 + - 7 | 3 | 2 | @ 24 ; o | 2 | ° N 4 oe) ro} 20% | Oo 25 D 2 4 0 | * 50a on WL | of ° Og a @ 6 0s ° 4 4 o, F 5 8% sm a | eee. @ 2 | 7 Rogen 4 #-3 -2 -1 0 1 2 3 4 Factor 1 (shape) Figure 6. Factor analysis (FA) plot of 180 Gastrocopta speci- mens from the Delmarva Peninsula of the Ll mixed samples determined to contain both forms. T. A. Pearce, M. C. Fields, K. Kurita, 2007 >I jee) Page areas of intermediate wetness, further supporting the idea that the two forms are distinct species. The dataset from Vanatta and Pilsbry (1906) and ours from Delmarva showed similar results, strengthening the conclusion that two species exist. The Delmarva dataset was more geographically restricted, but had more speci- mens, while the Vanatta and Pilsbry dataset was geo- graphically broader with fewer a cimens. Sources ee all Gastroc opta tappaniana included by Vanatta and Pilsbry (1906) are from locations more concentrated in NE USA, from Washington, DC to Maine ( (excepting one G. tap- paniana from Arizona), whereas their G. pentodon were more W idespread, from Texas to Iowa and Florida to Maine. This congruence despite different geographic sampling suggests that shell morphology does not vary significantly across geography. Bimodality of morphology supports the hypothesis of two species on Delmarva and throughout the eastern USA. Gastrocopta tappaniana on Delmarva occurred in samples from moister areas, whereas G. pentodon oc- curred in samples from a broad range of wetness. Sample wetness was unimodal and continuous, both for all samples and for samples containing at least 5 specimens, such that a difference in habitat by species does not seem to be an artifact of sample choice. This result confirms reports of Sterki (1906) and Pilsbry (1948) that G. tap- paniana occurs in moister areas. However, in contrast to authors who characterized G. pentodon as being in drier areas, our results agree with the report of Hubricht (1985) that G. pentodon has a wider moisture range and can overlap in habitat wetness with G. tappaniana. Geographically, G. tappaniana tended to occur on the SE part of the peninsula, and closer to water bodies, whereas G. pentodon was more evenly distributed across the peninsula, including its central ine Moistness seems to be an important ‘afinenise on the distribution of G. guia but some other aspect of geography might also play a role in its distribution, considering that fhe Atlantic coast samples from the SE part of the peninsula were the moistest samples. We examined the type specimen of Gastrocopta tap- 80 60 40 Individuals 20 -12 to -10 I paniana (Museum of Comparative Zoology, Harvard, lectotype MCZ 186171, paratype [not seen] 186172) but i re unable to locate type material of G. pe ntodon. For ,. tappaniana, Pilsbry (1948: S89) stated “Type locality, oo coll. Amherst College”, implying that an un- specified kind of type or types was present at Amherst. Bequaert and Miller (1973) observed that the holotype [sic] was transferred from Amherst to MCZ, mistakenly stating that Pilsbry (1948) referred to it as a holotype. C lench( 1965) noted that the type locality for G. tappa- niana is Roscoe, Coshocton Co., Ohio, and not Vermont as some writers have assumed. He chose a lectotype for G. tappaniana: “G. tappaniana appears to be a synonym of G. pentodon (Say); the lectotype (here selected) is nearest to the figure of pentodon given on pl. 3, fig. 7 1916, Manual oF Conch. (2) 24: 33, and not io fig. 9 which is given as tappaniana. This same plate was re- published i in Land Mollusca of North America, vol. 2, pt. 2, fig. 477, p. S87, Mono. no. 3, Acad. Nat. Sci. Phila- delphia, 1948.” Ve find it peculiar that Clench intended to choose a lectotype specimen of Gastrocopta tappaniana that was most like G. pentodon. Interestingly, despite Clench’s bias in his choice, the lectotype is consistent with G. tappaniana per both of our tests: it is located among the G. tappaniana specimens on factor plots, and it Glascihed as such by the DF derived in this paper. Vanatta and Pilsbry (1906) did not reveal their objec- tive criteria for classifyi ing the two forms, so researchers using their paper must accept their two groups at face vale. We recognize that DFA, by its nature, will find differences between nearly any groups. Therefore, to test whether the groups defined by Vanatta and Pilsbry (1906) are different from randomly selected eee we compared the percent correct classification of the DFA results for the groups as recognized by Vanatta and Pils- bry, to 10 randomized datasets with 41 specimens in one group and 12 in the other. In contrast to the 94.3% correct classification of groups as they defined, the randomized data sets averaged 70.9% correct classification. The higher correct classification of the non-randomized groups gives us Midry G. tappaniana med A wet 0 to2 nal 2to4 L j ato6 ba i 8 to 10 10 to 12 Discriminant Function Score Figure 7. classes. Histogram showing the number of individuals by discriminant function (DF) score, separately for each of three wetness Page 74 THE NAUTILUS, Vol. 121, No. 2 Figure 8. samples of Gastrocopta pentodon (open circles) and G. tappa- niana (solid circles). Symbols on the map reflect majority rule (24 samples contained both G. pentodon and G. tappaniana; the 5 samples with equal numbers of the two species are omitted). Delmarva Peninsula showing locations of 130 confidence that the two groups as defined by Vanatta and Pilsbry are non-randoin ( (p < 0.000001). Some disagreement exists in the literature regarding the correct classification of Gastrocopta carnegici Sterki, 1916. Sterki (1916a) described the new species as G. minuta, but subsequently changed the name to G, carnegiet because G. minuta was preoccupied (Sterki, 1916b). Sterki 1916a) did not designate a holotype among the three specimens, although Pilsbry (1948: S90, 892, fig. 450: 5) designated a lectotype by referring to the only unbroken specimen as the type. Turge -on et al. (1998) listed G. car- negiei as a valid species. Sterki (1916a) had indicated that G. carnegici is similar to G. tappaniana, but Hubricht (1985) considered G. carnegie to be a synonym of G. pen- todon. The DF, when applied to measurements taken from the lectotype illustration of G. carnegiei presented by Pilsbry (1948: 892, fig. 480, image | 5), classified the speci- men as G. pentodon uid the position it occupies in Fig- ure 3 (indicated by the plus sy seer suggest it is @losete to G. pentodon. The squat shape of ake lectotype figure is like G. tappaniana, with its large body whorl, bak it differs by a smaller shell and only 5 apertural teeth; the few whorls suggest that it might be an abnormal speci- men. Although we were unable to locate the lectotype of G. carnegiei at Carnegie Museum of Natural History, we located the two paralectotypes, which are broken, as noted by Pilsbry (1948), so their relevant shape measurements cannot be discerned. This study suggests that G. carnegiei falls within the range of variation of G. pentodon. Although separating the two species visually can be difficult, some characteristics might be helpful at distin- guishing between them. Gastrocopta tappaniana tended to be larger (wider shell, wider penultimate whorl width, and taller body whorl) and regarding shape, tended to have a shorter hat (section doov e the penultimate whorl) relative to both shell height and spire height (hat/h, hat/ spr), and a relatively taller body whorl. Regarding num- ber of teeth, Vanatta and Pilsbry (1906) stated that all G. tappaniana have 7 teeth (excluding teeth on the parietal wall), whereas G. pentodon have 5-9 teeth. However, according to their eainee the number of teeth for their G. pentodon ranged from 2-9. In contrast to their sug- gestion that tooth number might be a useful distinguish- ing character, our analysis suggested that number of teeth was not useful for separating species on the Del- marva Peninsula. A character that could be evaluated in future studies is whether the lower palatal fold of G. tappaniana is “usually not so long and entering as in G. pentodon” (Vanatta and Pilsbry, 1906; Pilsbry, 1945). Future work could aes molecular data such as DNA sequences to verify these conclusions. Another av- enue to address environmental influence on morphology would be to raise sibling specimens under different en- vironmental conditions. ACKNOWLEDGMENTS The U.S. National Science Foundation (DEB 9972026) funded part of this project. K. Fewlass Kling and others helped collect Delmarva samples and A. W. Doolittle helped with fieldwork. A. S. Italia, L. Brink Beebe, D. L. Scott, A. Gathers, and others diligently picked snails from leaf litter samples. We are grateful to Adam J, Baldinger, MCZ, for loan of the lectotype of Gastrocopta tappaniana. We are grateful to Amanda E. Zimmerman for expertise in producing the figures. Reviews by Jeff C, Nekola and an anonymous reviewer contributed substan- tially to improving this paper. LITERATURE CITED Bequaert, J.C. and W. B. Miller. 1973. The mollusks of the arid southwest, with an Arizona checklist. University of Arizona Press, ‘Tucson, xvi + 271 pp. T. A. Pearce, M. C. Fields, K. Kurita, 2007 Page 75 Clench, W. J. 1965. Note on Gastrocopta tappaniana (C. B. Adams). The Nautilus 78: 106—107. Goodfriend, G. A. 1986. Variation in land-snail shell form and size and its causes: a review. Systematic Zoology 35; 204— 233. Hagiuda, N. and K. Shigemasu. 1996. Junjo tsuki kategorikaru deta e no inshibunseki no tekiyo ni kansuru ikutsuka no chui ten. [Some remarks on the application of factor anayl- sis to ordered categorical data]. The Japanese Journal of Psychology 67: 1 1S. Hair, J. F., Jr., R. E. Anderson, R. L. Tatham, and W. C. Black. 1998. Multivariate data analysis, 5'" ed. Prentice Hall, 768 pp: Hubricht, L. 1976. Notes on some land snails of the eastern United States. The Nautilus 90: 104—107. Hubricht, L. 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana, Zoology, new se- ries 24: i-viii + 1-191 pp. Lauriol, B., E. Deschamps, L. Carrier, W. Grimm, R. Morlan, and B. Talon. 2003. Cave infill and associated biotic re- mains as indicators of Holocene environment in Gatineau Park (Quebec, Canada). Canadian Journal of Earth Sci- ences 40: 7S9-S03. Mardia, K. V. 1971. The effect of non-normality on some mul- tivariate tests and robustness to non-normality in the linear model. Biometrika 58: 105-121. Minton, R. L. and R. W. Gunderson. 2001. Puperita tristis (d Orbigny, 1842) (Gastropoda: Neritidae) is an ecotype of Puperita pupa (Linnaeus, 1767). American Malacological Bulletin 16: 13-20. Nekola, J. C. 2004. Terrestrial gastropod fauna of northeastern Wisconsin and the southern Upper Peninsula of Michigan. American Malacological Bulletin 18: 21-44. Nekola, J. C. and M. Barthel. 2002. Morphometric analysis of Carychium exile and Carychium exiguum in the Great Lakes Region of North America. Journal of Conchology 37: 515-531, Pearce, T. A. and A. S. Italia. 2002. Land snails and slugs in Delaware, USA: systematic survey reveals new distribution records. Western Society of Malacologists, Annual Report (for 2000) 33; 26. Pilsbry, H. A. 1948. Land Mollusca of North America (north of Mexico). Academy of Natural Sciences of Philadelphia, Monograph 3, 2(2): i-xlvii + 521-1113. Sterki, V. 1906. Additional notes on Bifidaria pentodon and B tappaniana. The Nautilus 19: 134-135 Sterki, V. 1916a. Description of a new Bifidaria. The Nautilus 29: 105. Sterki, V. 1916b. Note on Bifidaria minuta St. The Nautilus 30: S84. Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. Coan, F. G. Hochberg, W. G. Lyons, P. M. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A. Schel- tema, F. G. Thompson, M. Vecchione, and J. D. Williams. 1998. Common and Scientific Names of Aquatic Inverte- brates from the United States and Canada. Mollusks, 2nd edition. American Fisheries Society Special Publication 26, Bethesda, 526 pp. USDA-NRCS. 2004. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, Louisiana 70874-4490, USA. Consulted on 28 Feb. 2007. Vanatta, E. G. and H. A. Pilsbry. 1906. On Bifidaria pentodon and its allies. The Nautilus 19: 121-128. THE NAUTILUS 121(2):76-89, 2007 Page 76 Upper Jurassic Pleurotomariidae (Gastropoda) from southwestern Madagascar Steffen Kiel Earth Sciences M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution P.O. Box 37012 Washington, DC 20013- harasewych@si.edu Leeds LS2 9JT 7012 USA and University of Leeds UNITED KINGDOM Department of Paleobiology National Museum of Natural History Smithsonian Institution Washington, DC 20013- kiels@si.edu 7012 USA ABSTRACT This paper describes four new species of Upper Jurassic Pleu- rotomariidae from southwestern Madagascar: Obornella thompsonorum, Bathrotomaria annejoffeae, Bathrotomaria be- detteae, and Leptomaria takahashii. In addition, the previously described Leptomaria texta Delpey, 1948, is reassigned to the genus Obornella. Comparison of this fauna with that of the geographically proximal Kutsch region of northwestern India reveals it to consist of representatives of wide-ranging Tethyan genera, but also to exhibit strong endemism at the species level. INTRODUCTION A substantial number of well preserved Upper Jurassic pleurotomariid gastropods have recently been uncovered as a byproduct of commercial mining for ammonites in southwestern Madagascar. Six specimens, representing four species, were leadly made available to us for study by Mr. Chris 1 ‘akahashi. The pleurotomariids and am- monites were dug by village rs from pit quarrie s near the town of Zakar ha, in southwe sstern Madagascar. A survey of the literature on the Mesozoic gastropod fauna of Madagascar (e.g., De Ipey y, 1948; Collignon, 1949: Collignon, 1959; Kiel, 2006) revealed the majority of pleurotomariid species known from Madagascar to be of Cretaceous age, with only a single species, Leptomaria texta Delpey, 1948, reported from Jurassic strata. Al- though the sample available to us is of limited size and stratigraphic range, it expands our insight into the Juras- : pleurotomariid fauna of Madagascar. Like the well ‘loc ‘umented Jurassic pleurotom: wad fauna of the Kutsch also spelled Kachc hh region of western India (Jaitley et al., 2000; Das, 2002; Das et al., 2005), the Madagascar fauna reveals Tethyan affinities at the generic level, yet exhibits endemism at the species level. Both faunas evolved an increasing endemism within the Indo- Madagascan faunal province that was formed as the Tethys Ocean widened between Laurasia and eastern Gondwana, and a seaway started to develop between East and West Gondwana in the latest Jurassic (Titho- nian) (Hay et al., 1999; Shome et al., 2004). The scope of the present study is to review the Oxfor- dian (Upper Jurassic) pleurotomariid fauna of Madagas- car, describe five species, four of them new, and to re- view the relationships and biogeography of these pleu- rotomariids. GEOLOGICAL SETTING Rifting between Africa and Madagascar produced three large sedimentary basins along the west coast of Mada- gascar. These are, from North to South, the Ambilobe (or Diego), Mahajanga (also spelled Majunga), and Moron- dava basins. Sedimentation in this region commenced in the Carboniferous with the deposition of the Gondwanan Karoo sequences and equivalents. The first marine de- posits resulting from the break-up of the Gondwana su- percontinent are of Toarcian (late Lower Jurassic) age. From then on, alternating shallow marine, brackish, and fluviatile sediments were deposited in these basins. The pleurotomariids described here are from the Morondava basin, the southernmost of the three basins. Bio- and lithostratigraphic work in this basin is difficult because outcrops are few, index fossils are often not available, and measurable sections are usually short and difficult to cor- relate with each other. Consequently, only few litho- str: tigraphic units have been given formation names so far (Besairie and Collignon, 1972: I uger et al., 1994; Geiger and Schweigert, 2006), The quarries that produced the specimens described here are located to the west of the town Zakaraha (also M. G. Harasewych and S. Kiel, 2007 spelled Sakkara), one of them north of the Fiherenana River, the other to the south of it (Figures 1, 2). The fossiliferous lavers are thin and consist of pink-yellow iron-oolithic limestones, and are overlain by grey mud- stones (K. Bandel, pers. comm., 2007). The southern locality is very close, if not identical, with the “Amparam- bato section (VIb)” of Geiger and Schweigert (2006: 99) which was characterized by them as “a highly fossilifer- ous iron-oolitic limestone and sandstone bed with thin mud-stone interlayers.” Accordingly, the oolitic lime- stones contain an abundant ammonite fauna; the over- laying mudstones contain nodosariid Foraminifera and ostracods (Geiger and Schweigert, 2006). The northern quarry is geographically close to the “Middle-Upper Oxfordian Ankilimena section (XI)” of Geiger and Schweigert (2006). This section is characterized by re- current iron-oolitic limestones, which contain ammo- nites, rhynchonellids, bivalves, belemnites, echinoderms, and wood debris (Geiger and Schweigert, 2006: 103). Although Geiger and Schweigert (2006) did not report 4 A5et rs no? wane Figures 1,2. Location of collection sites. 1. Detailed map of two localities in southwestern Madagascar. 2. Location of sites on a map of land masses during the Late Jurassic. Page 77 (=) pleurotomariids or other gastropods from these sections, the remaining fossil content, their geographic position, and their lithology agrees well with that observed at the ammonite quarries visited by K. Bandel (pers. comm., 2007). The ammonite fauna of these two localities, especially the presence of Dhosaites cf. primus Collignon, 1959, suggests a ‘middle to upper Argovian (early Oxfordian)’ age (Collignon, 1959; H. Keupp, pers. comm., 2004). The term ‘Argovian’ has been abandoned due to its in- consistent usage, but largely falls within the range of the Oxfordian (Zeiss, 2003). The sediments considered as ‘Argovian’ by Besairie and Collignon (1972) correlate with those mapped as late Callovian-early Oxfordian by Geiger and Schweigert (2006). Thus, the pleurotomariids described here are most probably of Oxfordian (Upper Jurassic) age. SUPRASPECIFIC CLASSIFICATION WITHIN PLEUROTOMARIIDAE The number of genera and subgenera currently recog- nized within the family Pleurotomariidae (Appendix 1) has nearly doubled since the family was reviewed in the Treatise of Invertebrate Zoology (Knight et al., 1960). Of the 21 genera and subgenera now recognized (Figure 3), 16 are Mesozoic. Of these, five are restricted to the Tri- assic, three to the Jurassic, and only a single subgenus to the Cretaceous. At present, Leptomaria and Conoto- maria are the only genera that are recognized as having survived from the Mesozoic into the Cenozoic. Of the seven Cenozoic genera, four survive in the Recent fauna. According to the literature surveyed, there is no chrono- logical overlap between the Mesozoic genera and the Cenozoic genera. As noted by a number of researchers (e.g., Hickman, 1976; 1094: Szab6, 1980: 49; Das, 2002: 99) fossil pleu- rotomariids are difficult to classify objectively, since the criteria upon which fossil pleurotomariid genera are de- fined differ conspicuously from those applied to Ceno- zoic genera. The monophyly and phylogenetic relation- ships of the surviving Cenozoic genera have been con- firmed using molecular data from living representatives (e.g., Harasewych et al, 1997; Harasewych, 2002). By contrast, the relationships of Mesozoic genera and the species assigned to them are far less certain, as generic classification tends to be based on relatively few con- spicuous morphological features (Table 1) especially those that are most easily derived from poorly preserved specimens and external or internal molds, while other characters are unconstrained and may vary widely. Szabé (1980: 49) commented that “almost all genera can be identified with certainty” on the basis of the shape of the whorl section and the surface of the whorls, as well as the position and width of the selenizone. Conti and Szabd (1987: 43) raised a question as to the significance of the presence or absence of an umbilicus in pleurotomariid Page 78 THE NAUTILUS, Vol. 121, No. 2 MESOZOIC TRIASSIC JURASSIC |CRETACEOUS U © @ ° OQ io) a © Figure 3. Geological ranges of the {¢ ssil rect ord classification. The presence or absence of an umbilicus is sufficient to distinguish the most basal dichotomy among living Pleurotomariidae, yet this feature remains uncon- strained and may vary widely within most Mesozoic gen- era and even within some “species” as they are currently diagnosed. It is therefore not surprising that fossil species are frequently reassigned from one genus to another e.g., Pyrgotrochus to Laevitomaria Conti and Szabo, LOST 2005 16: Perotrochus to Leptomaria, see Das et al., 33] served specimens become available especially as more numerous and better pre- ausd0bIlO genera and subgenera within the family Pleurotomariidae, CENOZOIC QUad0I|4 QU800}SI9|q GENERA OF PLEUROTOMARIIDAE Mamoeatomaria Begg and Grant-Mackie, 2003 Tahua Begg and Grant-Mackie, 2003 Murihikua Begg and Grant-Mackie, 2003 Pleurotomaria Defrance, 1826 Ornatospira Pan, 1982 Stuorella Kittl, 1891 Talantodiscus P. Fischer, 1885 Pyrgotrochus P. Fischer, 1885 Anodotomaria Szabo, 1980 Bathrotomaria Cox, 1956 Cyclostomaria Szabo, 1980 Laevitomaria Conti and Szabo, 1987 Obornella Cox, 1959 Conotomaria Cox, 1959 Leptomaria E. Eudes-Deslongchamps, 1864 Indomaria Das, 2002 Chelotia Bayle in P. Fischer, 1885 Entemnotrochus P. Fischer, 1885 Perotrochus P. Fischer, 1885 Bayerotrochus Harasewych, 2002 Mikadotrochus Lindholm, 1927 arranged by first occurrence in the SYSTEMATICS Family Pleurotomariidae Swainson, 1S40 Genus Obornella Cox, 1959 Obornella Cox, 1959: 238. Type Species: Pleurotomaria plicopunctata J. é Eudes-Deslongchamps, 1849 (By original designe ney Bajocian (Middle Jurassic) of France. Diagnosis: The genus Obornella is characterized by a shell that is low turbiniform to sublenticular, with a nar- Harasewych and S. Kiel, 200 = x. — = spray pur MOTeYS yuasqe XOAUO. [LoyAr-prut sp109 yeadg ‘peotg 10 YRoAy A[BUO.QS sugqnqy yuosqy yuosqy yuasqy QAOqY — popls-P promos snyoo.jo1ahivg sp.l09 MOTeYS quasqe [LOYA\-plUt jeuds osopon ‘proig, 10 yRaAy XOAUOD) Suyuqy quasqy yuosqy juosqy MOLIG — Papls-p fRolloy — smpoo.gopvyiyy asopou 10 out MO]EYS quasqr [LoTp\-plul spi09 peurdg “proig IO YROA\ XOAUOD) suyqqnqy quasqy quasqy yuosqy Mopaq ysuf— papts-F jeowog SM IOMOMIT spioo pur doop yuasqe [Loya-prut sprampy pemdg = ‘MoumN 10. YRoA\ rl ysnya opU yuasqy yuosqy QA0qY — popls-F [ROOD SNYIOAJOULWAPUTT sptoo daap quasqe [HOYA\-prutt jetids asopoN =“ MOLIBN LO. YROAY Wi ysty a opU yuosqy yuasqy QAOGY — papls-f [Rolulon pyopayy aqe]jeours MOTEYS yuasqe x apr 0} sptoo prudg = ‘Mourn 10. YRay A[BUO.S Suyquqy quasqy juasqy uosqy [LOY A-plUL PY Popts-F [Rowwo9g pupwojday doop Opi 0} [toyar-prut spi0o yeudg = ‘more Usa wll sup] yuosqy juosqy yuosqy aAoqe 10 FY papts-c peowwod DLIDWOJOUDT URUTULOp avysoo MOTPYS juosqr. XOAUOK ‘oPRTPOOURD, = SMOLIBNY LO ROA, A[BUOLYS SUNY Asa. Juosa.tg quesqy = Atoyditod uwon papts-p [row DIJOULOGO yuasqe [LoyA\-prut sp0o peardg MOLE IO YROA\ Wa possordury opi yuasqy yuasqy MOOG — Popls-F jeomtoy DUDUONAIVD'T] spear peatids asopou MO.LIRU Atoydiiod ‘avysoo [RIXy MO.LIRN \uaso.g Rl ysnyy 0 Juasqy UIs yuasqy aaoqe ysuf papts-F [Rolo DIJQLON]S spurq asopou asopou MO.LIVU juosoid [LOYPA\-piut ‘spa09 ped prog \uasotg ra ysnpy 10 yuasqy yuasotg SOpON MOLI — popls-p [Romo smyoospod.thg }USUTULOp peixe ‘oqyyyjoourg, MO.LIEN quasqy Wp Apron sunynqy op quasqy yuasqy [LOYA\-PIU PY papis-p — [ROLWOD AO] v.ndsoppuUui 0.8 aperture length), ovate, thin, corneous, with terminal nucleus. Radula (Figures 7-9) short (0.33 aperture length) con- sisting of 45-51 rows of teeth (7 per row). Rachidian teeth broad, with wide, curved basal plate, with strong central cusp flanked by 5-8 short, conical denticles. Lat- eral teeth broad, with stout bases, 6-8 denticles along ventral edge. Marginal teeth scythe-like, with eaieath edges. Jaws (Figure 10) paired, narrow dorsally, ex- panded ventro-laterally, with fringed edges. Type Locality: Oahu, off the north shore district of Haleiwa, muddy sand, in red shrimp traps set at 300- 390 m. Type Material: Holotype, USNM_ 1099759; Paratype 3, USNM 1099760, Paratypes 1, 2, 4, Takahashi collec- tion. All from the type locality. Additional Material Examined: AUSTRAL ISLANDS: MNHN, Raevavae, 23°50.54 S, 147°42.73° W, in 400 m. BENTHAUS: stn DW 1884, Iles are 570-620 m (1 dd): sth DW 1885, as last, 700—SO0 m (1 dd); stn DW 1897, ouest de Rapa, 480-700 m (2 dd); stn DW 1899, as last, 342-600 m (1 dd); stn DW 1903, Banc NE Rapa, 400-600 m (1 dd); stn DW 1923, Recif Nielsen, 360-840 m (1 dd): sth DW 1924, as last, 340-800 m (1 dd): stn DW 1925, as last, 560-790 m (1 dd): sth DW 1929, Banc Président Thiers, 350-370 m (1 dd); stn DW 1932, as last, 500-600 m (1 dd): stn DW 1933, as last, 500-859 m (2 dd); stn DW 1940, N de Rae vavae, LOO—460 m (3 dd); stn DW 1943, as last, 950 m (2 dd): sth DW 1945, Banc Lotus, 120-500 m (2 dd); stn DW 1951, as 206-450 m (1 dd): stn DW 1957, Tubuai, 55S—1000 m (2 dd): stn DW 1961, as last, 470-800 m (3 dd): stn DW 1973, Banc Arago, 300-350 m (2 dd); stn DW 1974, as last, 450-615 m (2 dd): stn DW 1992, Rurutu: Mont de Lotus, 442-444 m (1 lv): stns 1997-2001, Rurutu, 200—LOOO m (2 dd); stn DW 1998, cote N de Rurutu, 250-302 m (1 dd?); stn DW 1999, as last, 270-500 m (4 dd); stn DW 2000, cote N de Rurutu, 270-480 m (1 dd); stn DW 2001, port de Rurutu, 200-550 m (1 Iv?); stn DW 2006, cote E de Rurutu, 35-450 m (1 dd); stn CAS 2008, cote E de Ru- rutu, 280-300 m (2 dd); sth DW 2018, Rimatara, 770— 771 m (1 dd); stn DW 2021, Rimatara, 1200-1226 m (1 dd). Distribution: Sassia melpangi is broadly distributed throughout the central West Pacific, from Guam to Ha- waii and Raevavae in the Austral Islands, but does not occur in the Marquesas. [t inhabits upper bathyal depths, with live specimens collected between 200 and 550 m. Etymology: This species is named in honor of Mr. Melvin Pang, of Oahu, Hawaii, who collected the type series. Comparative Remarks: ~The new species Sassia mel- pangi is readily distinguished from the western Pacific S. M. G. Harasewych and A. G. Beu, 2007 Page 93 Figures 7-13. Sassid ™meé lpangi new species of radula near distal end. lateral teeth spread to reveal rac hidian teeth. 9. Right lateral view or radular ribbon. Scale bar s 7-9). 10. Jaw of holotype, with detail of edge. 11. Periostracum of paratype 1, at midpoint of final vari 13. Lateral views of protoconch of paratype 1. Scale bar | mm (applies to Figures 12, 13 7-9. Radula of the holotype. 7. Dorsal view of radula near mid-length. 8. Dorsal vic LOO jxn 12. Apical | applies to Figur Page 94 THE NAUTILUS, Vol. 121, No. 2 remensa and 8S, semitorta in having: more evenly rounded whorls that lack a pronounced shoulder the presence of more numerous, evenly spaced axial ribs that form a reticulate surface sculpture; a more evenly ovate aper- ture surrounded by an inductura with ventrally reflected edges; as well as a strong pigmentation pattern along the outer lip. Sassia melpangi most closely resembles S. nas- sariformis from southeastern Africa, with which it shares the rounded whorl profile and even cancellate sculpture. Sassia nassariformis can be differentiated by its shoul- dered varices, smaller aperture, weaker pigmentation along the outer lip, and by its thicker inductura, which is more triangular than ovate, and which is not reflected along it edges. Sassia melpangi has a broad range throughout the tropical Pacific. While the type lodalitay is off the Island of Oahu, a number of specimens are known from French Polynesia, and photographs of two specimens from Guam, Marianas Islands have been examined. This spe- cies inhabits upper bathyal depths (300-400 m). The position of worm tubes (Figure 5, wt) on the ventral surface of living specimen suggests that Sassia melpangi is epifaunal atid inhabits hard “Sabsirates. ACKNOWLEDGMENTS We are grateful to Chris Takahashi, for bringing this material to our attention, and for donating the holotype and one agg to the National Museum of Natural History. Richard Salisbury kindly made available photogr ae of two specimens dredged off Guam, Mari- anas Islands, and Professor Alison Kay provided photo- graphs of additional specimens dredged in Hawaii. LITERATURE CITED Beu, A. G. 199Sa. Superfamily Tonnoidea. In: Beesley, P. L., G. J. B., Ross, and A. Wells (eds.) Mollusca: The Southern Synthesis. Fauna of Australia 5. CSIRO Publishing, pp. 792-803. Beu, A. G. 1998b. Indo-West Pacific Ranellidae, Bursidae and Personiidae (Mollusca: € Gastropod: 1). A monograph of the New Caledonian fauna and revisions of re lated taxa. Mem- oires du muséum national dhistoire naturelle 178: 255 pp. Cossmann, M. 1903. Essais de Paléoconchologie comparée. Vol. 5. M. Cossmann, Paris, 215 pp. Habe, T. 1961. Coloured Hlustrations of the Shells of Japan vol. 2. Hoikusha Publishing Co., Osaka, ix + 182 pp, appendix 42 pp. Iredale, T. 1936. Australian molluscan notes. No.2. Records of the Australian Museum 19: 267-340. Sowerby, G. B. HI, 1902. Mollusca of South Africa. Marine Investigations in South Africa 2: 93-100. Watson, R. B. 1881. Mollusca of the HMS Challenger Expe- dition, part 7. Families Pyramidellidae, Naticidae, Cassidae, Tritonidae. Journal of the Linnean Society of London 15: 245-274. THE NAUTILUS 121(2):95-98, 2007 Page 95 A new species of Microcancilla (Gastropoda: Cancellariidae) from the continental slope off northeastern Brazil José Carlos N. de Barros Departamento de Pesca e Aquicultura Richard E. Petit S06 St. Charles Road Universidade Federal Rural de Pernambuco North Myrtle Beach, SC 29592 USA Avenida Dom Manuel de Medeiros, S/N, Dois Irmaos 52171-030, Recife, BRAZIL mundovan4t@yahoo.com.br r.e.petit@worldnet.att.net ABSTRACT Microcancilla jonasi new species is described from deep waters off northeastern Brazil. The genus Microcancilla Dall, 1924, had not previously been recorded in Brazilian waters. Speci- mens were collected from sediment dredged as part of the REVIZEE program (Assessment of the Sustainable Potential of Living Resources in the Exclusive Economic Zone) during prospecting work on the continental slope off the state of Per- nambuco at depths of 425 to 690 meters. The present study reveals that these specimens present affinities to the species Microcancilla microscopica (Dall, 1889), which differs from the new species, among other features, by having strong spiral sculpture between sigmoid axial ribs. In addition, a lectotype for Cancellaria microscopica Dall, 1889, is designated herein. Additional keywords: Cancellaria, Neogastropoda, Pernam- buco. INTRODUCTION In his work on the gastropods collected in the West In- dies (1879-80) by the U.S. Coast Survey Steamer BLAKE, Dall (1889a) described Cancellaria microscopica Dall, 1889, based entirely on empty shells. Dall later (1889b: 106) placed that species with a ‘toa in the genus Ad- mete ie er in Moller, 1842. Still later, Dall erected the cancellariid genus Microcancilla Dall, 1924. When intro- ducing this genus, Dall did not give a genus description, and listed only the single species Admete [sic] micro- scopica Dall, 1889. Until now no congeners have been reported. Most species of Admete are found in shallow waters of polar regions but a few occur in deepe r water in tem- perate zones (Harasewych and Petit, 1986; Knudsen, 1964). However, all of the taxa described as Admete have not been studied in detail and it is probable that some are not properly placed in this genus (Bouchet and Warén, 1985). A recent study of deep-water Cancellaridae from the New Caledonia area (Bouchet and Petit, in preparation) shows that the central Pacific fauna contains species clearly attributable to Microcancilla and others that are morphol ogically similar to Admete aethiopica Thiele, 1925, font Somalia [illustrated in Verhecken (1997: 312, fig. 52)|. Intermediate between these seemingly dispar- ate morphologies and having various features in common with them are a series of species. Verhecken (1997), figuring the Somalia specimen, placed it in the genus Adee with a query but offered no other possible plac e- ment. It is not contended here that Microcancilla jonasi new species, “Admete” aethiopica Thiele, and Microcancilla microscopica (Dall) are truly congeneric species. How- ever, Microcancilla is considered at the moment the best available placement within the existing genera of the Cancellarioidea and such placement is provisional. Until the small deep-water species of Cancellariidae are better understood and the validity and limits of other available genus group taxa are established, we do not wish to in- troduce a new genus-group taxon. This problem with the available genera for small cancellariids was succinctly pointed out by Maxwell (1992: 167). Microc ancilla jonasi new species is considered endemic to the continental slope off the state of Pemambuco, Brazil. MATERIALS AND METHODS All specimens examined were obtained in 1999-2000, during oceanographic prospecting work on the continen- tal slope off the state of Pernambuco, Brazil. No live specimens were collected. Shells were mounted on specimen stubs and exmained and photographed under a Jeol JSM 6360 Scanning Electron Microscope, at the “Instituto Tecnolégico de Pernambuco (ITEP)”. Shells were measured using a stereo microscope with eyepiece micrometer. The type material was deposited at the Academy Natural of Sciences, Philadelphia, USA (ANSP): Museu Nacional, Rio de Janeiro, Brazil (MNRJ); Museu de Zoologia da Universidade de Sao Paulo, Brazil (MZUSP):; Museu Oceanogratico do Rio Grande, Rio Grande, Brazil (MORG). Page 96 THE NAUTILUS, Vol. 121, No. 2 SYSTEMATICS Family Cancellariidae Forbes and Hanley, 1851 Genus Microcancilla Dall, 1924 Type Species: © Admete microscopica (Dall, 1889) [= Cancellaria microscopica Dall, 1889a] by original desig- nation. Recent, Caribbean. Microcancilla jonasi new species (Figures 1-8) Description: — Shell conical, small, fragile, translucent, whitish, short spire. Protoconch smooth, globose, pau- cispiral with 1.5 whorls, terminating with the emergence Figures 1-8. Scanning electron micrographs of Microcancilla jonasi new species (all deposited in MOR¢ of the axial sculpture of the teleoconch (Figures 7-8). Teleoconch with 2.5 gee convex whorls. Suture well- marked, with a small, flattened subsutural region orna- mented by the adapical portion of the axial “ibs. On the margin of this region, there is a strongly nodular shoul- der, nodule 4S coinciding with axial ribs below, resembling a small crown. Below the shoulder, there is a second weaker spiral cord, with nodules somewhat smaller than those of the crown on margin of a small, concave, spiral depression (Figure 6). Body whorl developed, very wide, with about 69% of the total size of the shell, sculptured with 15 to 18 rounded, regularly spaced ribs, which dis- appear toward the base. Interspaces ornamented by ob- scure threads. Base imperforate, strongly conical, with +): L. Holotype, apertural iew, leneth fmm; 2. Paratype, apertural view, length: 3.28 mm; 3. Paratype, apertural view, length = 3.12 mm; 4. Paratype ipertural view, length = 2.55 mm; 5-8. Paratype, length: 3.60 mm Apertural view. 6. a view, showing strongly nodular shoulder and weak spiral cords (Figure 5). 7-8. Protoconch (shell in Figure 5). Scale bars: Figure 6 = 500 jm; Figure 7 = 200 jzm; Figure 8 = LOOwm J.C. N. de Barros and R. E. Petit, 2007 Page 97 convex contour, ommamented by 3 to 5 weak spiral cords that emerge from the interior of aperture. Aperture el- liptic. Outer lip fragile and smooth inside. Inner lip cad y reflected, wide parietal region, without callus, median portion of the columella with reflected appear- ance, thickened upon the umbilical wall with no columel- lar folds. Final anterior portion of the columella inclined to the left of the shell axis. Type Material: —© Holotype, MORG 50.716, length = 4 mm; 4 paratypes, MORG 50.718 (Figures 2-8); 3 paratypes, MZUSP $1755; 3 Paratypes ANSP 413549, 1 paratype, MNRJ 1OS39, Pernambuco, Brazil, D-4, 08°42.1" S, 34°44.1' W, 425 m, muddy bottom, 25 Mar. 2000; 2 paratypes, MNRJ 10835; 3 paratypes, MZSP $1756; 1 paratype MORG 50.717 (Pernambuco, Brazil, D-11, 08°46.5' S, 34°44.5' W, 690 m, muddy bottom, 18 Sep. 2000). Type Locality: Pernambuco, northeastern Brazil, D-4, 08°42.1' S, 34°44." W, 425 m, muddy bottom, 25 Mar. 2000. Etymology: The species is named after Mr. Rainer Jonas, scientific director of the Gesellschaft fiir Biotech- nologische Forschung (GBF), Germany, for his assis- tance in obtaining literature and his constant support during the identification work of gastropods from the contnenals slope of Brazil. Geographic Distribution: — Endemic to the Conti- nental Slope of the State of Pernambuco, northeastern Brazil. Habitat: © Muddy substrate, 425 to 690 m. Discussion: — Microcancilla jonasi new species studied here is distinguished from Microcancilla mic roscopica and ?Admete aethiopica based on the following charac- teristics: (1) sculpture of the body whorl: M. micro- scopica has a strong spiral sculpture, with subequal rather coarse threads, forming a reticulum with the axial spirals. 2A. aethiopica presents a wide body whorl, with a strong axial sculpture and smooth spiral bands se parated by narrow grooves disappearing near the base, M. jonasi has strong axial ribs, sinuous and rounded, and an ob- scure spiral ornamentation with no reticulation; (2) Spi- ral ornamentation: M. microscopica has threads that al- ternate between strong and weak, crossed by raised growth threads, ?A. aethiopica has strong axial ribs crossed by weaker spiral threads, M. jonasi only presents raised axial ribs: (3) Shoulder: on M. microscopica, the shoulder is obsolete and the subsutural platform be- comes rounded on the body whorl, ?A. aethiopica pre- sents an angular shoulder, strongly nodular, and a nar- row, flat subsutural platform, M. jonasi has a small, flat subsutural platform, bordered by a nodular shoulder with a lower depression and followed by a second sub- shoulder cord that is more weakly nodular: (4) Umbili- cus: M. microscopica has a distinct, moderate umbilicus with no bounding carina or siphonal fasciole, 2A. aethi- opica and M. jonasi do not possess an umbilicus; (5 Continental Slope from the State of Aperture: M. microscopica has an aperture that is rounded behind and strongly angular in front, ?A. aethi- opica has an oval aperture, slightly square-cut adapically, M. jonasi has an elliptic aperture, weakly constricted be- hind: (6) Inner lip: M. microscopica has a moderately callous inner lip with one extremely faint fold about the middle, PA. aethiopica has an inner lip with two very weak folds near the halfway height, M. jonasi has a re- flected inner lip, wide parietal region, with no folds: (7 Columella: In M. microscopica and PA. aethiopica, i columella is straight, parallel to shell axis; in M. jonasi, it is inclined to the left of the shell axis. Only the type species has been allocated in Microcan- cilla since the genus was ae The specimen fig- ured by Dall (1902: pl. 29, fig. 4; 1903: pl. 75, fig. 4 [same Figures 9-11. Scanning electron micrographs of Microcan- cilla microscopica (Di ull, 1889 lectotype, USNM 82977. 9. Apertural view. 10. Lateral view. 11. Protoconch. Scale bars Figures 9, 10 = 1 mm; Figure 11 = 200 pm Page 98 THE NAUTILUS, Vol. 121, No. 2 drawing]) and illustrated herein (Figures 9-11) is more rounded at the anterior than in most specimens in the syntype series and also has a heavier columellar callus. Kaicher (1978: Card 1940) Pee al illustrated this same specimen from the USNM type collection (USNM 62977 |sic; error for Som as Admete micro- scopica (Dall), Admete being the genus used by Dall in 1889b and 1903. She incorrectly felenied to this speci- men as holotype but Dall did not designate a type speci- men. The species was originally deserbe d from two lo- calities and there are numerous specimens in the original lots. Under Article 74.5 of the current Code (Interna- tional Commission on Zoological Nomenclature, 1999) Kaicher’s statement does not “qualify as a lectotype des- ignation. In order to rectify this, USNM 82977, off Yucatan, 366 m (originally 200 fathoms), is here desig- nated lectotype of Cancellaria microscopica Dall, 1889. The type locality thus becomes restricted to Campeche Bank, off Yucatan, Mexico. ACKNOWLEDGMENTS We are grateful to Mr, Enilson Cabral of the Research and Management Center of Fishing Resources of the Northeastern Coast — CEPENE/IBAMA for the collec- tion of the material analyzed in the present work. This study Was par tially supported by the Assistance to Sci- ence and Technology Foundation of Pernambuco (FACEPE/CNPq). Dr. M. G. Harasewych, National Museum of Natural History, Smithsonian Institution, W ashington, DC, kindly furnished the SEM of Microcancilla microscopica (Dall) and reviewed the manuscript. LITERATURE CITED Bouchet, P. and Warén, A. 1985. Revision of the Northeast Atl ee bathyal and abyssal Neogastropoda excluding Tur- ridae (Mollusca, Gastropoda). Bollettino Mal: acologico, suppl. 1: 121-296. Dall, W. H. 1889a. Reports on the results of dredgings, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U. S. Coast Survey Steamer “Blake”, Lieut.- Com- mander ©. D. Sigsbee, U.S. N., and Commander J. R. 3artlett, U.S. N., commanding, XXIX. Bulletin of the Mu- seum of Comparative Zoology 1S: 1-492, pls. LO-40. Dall, W. H. 18S9b. A preliminary catalogue of the shell-bearing marine mollusks and brachiopods of the southeastern coast of the United States, with illustrations of many of the species United States National Museum Bulletin 37: 1-221, pls. 1-74. [Reprinted 1903 with additional plates: 1-232, pls. 1-95.]. Dall, W. H. 1902. Illustrations and descriptions of new, unfig- ured, or imperfectly known shells, chiefly in the U. S. National Museum. Proceedings of the United States Na- tional Museum 24(1264); 499-566, pls. 27-40. Dall, W. H. 1903. A preliminary catalogue of the shell-bearing marine mollusks and brachiopods of the southedertn coast of the United States, with illustrations of many of the species. United States National Museum Bulletin 37: —232, pls. 1-95. Dall, W. H. 1924. Notes on molluscan nomenclature, Proceed- ings of the Biological Society of Washington 37: S790. Harasewych, M. G. and Petit, R.E. 1986. Notes on the mor- phology of Admete seein (Gastropoda: Cancellariidae). The Nautilus 100(3); 85-91. International een on Zoological Nomenclature, 1999. International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, xxix + 306 pp. Kaicher, S$. D. 1978. Pack #19, Cancellariidae. of world-wide shells. Cards 1559-1964. Knudsen, J. 1964. Scaphopoda and Gastropoda from depths exceeding 6000 meters. Galathea Report 7: 125-136, Maxwell, P.A. 1992. Eocene Mollusca from the vicinity of Mc- Culloch’s Bridge, Waihao River, South Canterbury, New Zealand: Pale oecology and systematics. New Zealand Geo- logical Survey Pe leontological Bulletin 65: 1-280. Petit, R.E. and Harasewych, M.G. 2005. Catalogue of the su- perfamily Cancellarioidea Forbes and Hanley, 1851 (Gas- tropoda: Prosobranchia) - 2"4 edition. Zootaxa 1102: 1-161. Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-E — I. Teil. Deutsche Tiefsee-Expedition 1595-1899, 17(2 35-382, Pls. 13-46. Verhecken, A. 1997. Mollusca Gastropoda: Arafura Sea Can- cellariidae collected during the Karubar Cruise. Mémoires du Muséum National d'Histoire Naturelle 172: 295-323. Card catalogue THE NAUTILUS 121(2):99-103, 2007 Page 99 A new species of Gerdiella (Gastropoda: Cancellariidae) from the South Atlantic Ocean off Brazil with discussion of an undescribed species Silvio Felipe B. de Lima José Carlos N. de Barros Departamento de Pesca e Aquicultura Universidade Federal Rural de Pernambuco Avenida Dom Manuel de Medeiros, S/N, Dois Irmaos 52171-030, Recife, BRAZIL stblima@yahoo.com.br mundovan4@yahoo.com.br re. petit@att.net Richard E. Petit 806 St. Charles Road North Myrtle Beach, SC 29582-2846 USA ABSTRACT Two rare species of Cancellariidae were identified during the study of material from oceanographic dredge hauls indlepalken in 2000 by the fishing vessel NATUREZA in ‘deep waters off the state of Pernambuco, Brazil. The species belong to the genus Gerdiella Olsson and Bayer, 1972. Gerdiella alvesi new species is similar to Gerdiella c ‘ingulata Olsson and Bayer, 1972, as both have strong, nodular ornamentation that is coarsely cancellated and a heavily thickened, lirated outer lip. A second species, Gerdiella sp.. is identified based on the protoconch, cancellated ornamentation and the presence of two columellar folds, dis- tinguished from the species described herein by its ornamen- tation and the absence of a subsutural keel. Additional keywords: Mericella, Neogastropoda, bathyal, Per- nambuco INTRODUCTION The family Cancellariidae Forbes and Hanley, 1851, represented by a large number of fossil and recent gas- tropods da bated among diverse marine regions throughout the world. The group inhabits subtidal to bathy al sandy and muddy bottoms of tropical and tem- perate regions, with the greatest diversity found along the eastern Pacific coast of the Americas and the ceniel Indo-Pacific area (Harasewych and Petit, 1982). In the western Atlantic Ocean, the number of known species is still relatively small especially with regard to the Brazilian coast (see for instance Harasewych et al. 1992). The genus Gerdiella Olsson and Bayer, 1972, was in- troduced to include three species described by these two authors from bathyal depths of the Florida ‘Straits and south of Jamaica. These species are: Gerdiella gerda from the Straits of Florida, 648-622 m: G. santa from the Straits of Florida, 64S—622 m: and G. cingulata from S of Jamaica, 549-530 m. Another specimen of G. cingulata, collected in 1961 by R/V OrEGON, sta. 3552, 130 miles ESE of New Orleans, Louisiana, 29°07’ N, 88°05’ W, trawled in 732 m, is now catalogued as USNM 811462. No additional species of Gerdiella have been discovered until now. The genus Mericella Thiele, 1929, was introduced by Thiele to accommodate the bathyal Mericella jucunda (Thiele, 1925) from off Tanzania. He originally placed the species in Cancellaria (Merica). Morice. bozzetti Petit and Harasewych, 1993, was described from off So- malia. Petit and Harasewych at the same time placed Cancellaria (Merica) paschalis Thiele, 1925, in the genus Mericella. Mericella paschalis was described from a bro- ken fragment, but recently collected material from off of Mozambique allowed Verhecken and Bozzetti (2006: 15- 16) to confirm the allocation of the species in Mericella. In a recent paper, Verhecken and Bozzetti (2006) ae Gerdiella in the synonymy of Mericella Thiele, 1929. As observed by those two authors, Mericella was ed by Olsson and Bayer in the original description of Gerdiella. Verhecken and Bozzetti (2006: 17) stated that the two genera are “very much alike conchologically, the main differences being ‘the relative spire height and the suture form.” They also considered relative aperture heights, observing that, as Gerdiella has a shorter aper- ture, the ratio in this latter genus agrees “with Petit and Harasewych (1993: 223) Go comer a value of 50.5 a diagnostic feature for Mericella.” Verhecken and Bozzetti did not point out that Petit and Harasewych used additional characters to differentiate these genera. Verhecken and Bozzetti also stated that “there are no important differences in shell characteristics that would justify a separation between Mericella and Gerdiella.” Although shown on their table, the text does not mention the fact that Gerdiella species have axial ribs on the pro- toconch. However, in an earlier work Verhecken (ohos: Page 100 THE NAUTILUS, Vol. 121, No. 2 513) stated that “protoconch characters are not consid- ered of diagnostic importance at generic level by this writer.” We disagree with that approach, especially when protoconch characters allow for the distinction of west- ern Atlantic taxa from those from the Indian Ocean. We consider Gerdiella to be a valid genus with species known at present only from the w vestern and southern Atlantic Ocean. The geographical grouping of Mericella, with all known species being from off eastern Africa, and Gerdi- ella, with all known species being from the western and southern Atlantic, is obvious. Verhecken and Bozzetti (2006: 17) mentioned that the eastern Pacific Cancellaria corbicula Dall, 1908, was placed in Gerdiella by Kaicher (1978: card 1952). We do not agree with that placement as the species has a euneatl protoconch and an aperture height greater than one-half shell height. Its generic placement remains in doubt. Verhecken (2002: 512) studied three juvenile and frag- mented shells collected from the Continental Slope at Pernambuco, northeastern Brazil, during probes of the CHALLENGER EXPEDITION in 1873. Those specimens were considered by him to be conspecific and the pos- sibility that they represent a new species of Gerdiella was mentioned. We agree with Verhecken that more and better specimens are needed for identification to be con- firmed. In this paper we describe a fourth species of Gerdiella from the Western Atlantic, collected from the Continen- tal Slope off Pernambuco, Brazil. This is the first definite record of the genus for the South Atlantic. The soft parts and radula are as yet unknown, but the conchological characters are sufficient to justify the naming of a new species. MATERIALS AND METHODS All specimens examined were obtained by the research vessel NATUREZA along the Continental Slope off Per- nambuco during oceanic prospecting work for the Re- search and Management Center of Fishing Resources of the Northeastern Coast—CEPENE/IBAMA. At the Ma- lacology Laboratory of the Universidade Federal Rural de Pernambuco, the specimens were sorted under a ste- reomicroscope, cleaned in a diluted detergent solution, rinsed in distilled water, and air-dried. Shells were mea- sured using a stereomicroscope with eyepie ce microme- ter and photographed with a Nikon COOLPIX 885 digi- tal camera. Scanning electron micrographs were made using a Jeol JSM 6360 Scanning Electron Microscope at the Electron Microscope Laboratory of the “Instituto Tecnolégico de Pernambuco (ITEP)” Abbreviations used: ANSP, Academy of Natural Sci- ences Philadelphia: LMUFRPE, Laboratério de Mala- cologia da Universidade Fedral Rural de Pernambuco, Brazil; MNRJ, Museu Nacional, Rio de Janeiro, Brazil: MORG, Museu Oceanografico do Rio Grande, Rio Grande do Sul, Brazil; MZUSP, Museu de Zoologia da Universidade de Sao Paulo, Brazil. SYSTEMATICS Family Cancellariidae Forbes and Hanley, 1851 Genus Gerdiella Olsson and Bayer, 1972 Type Species: Gerdiella gerda Olsson and Bayer, 1972 by original designation. Recent, Caribbean. Gerdiella alvesi new species (Figures 1—5) Description: Shell fusiform, stout, white, strongly or- namented axially and spirally, entirely marked by growth lines between spiral cords. Protoconch globose, cap- shaped, with 1.5 whorls, ormamented by numerous mi- croscopic spiral threads and weak axial ribs on final por- tion. Transition to teleoconch marked by strong raised axial rib. Nucleus small when compared with rest tof pro- toconch, which is inflated. Teleoconch with 5.5 whorls. Spire narrow, smaller than body whorl. Whorls rounded, strongly omamented, with reticulated sculpture. Spiral cords intersect the equally strong axial ribs, forming strong nodules with a pustulose aspect, which progres- sively increase in size toward body whorl. First whorl with 20-22 axial ribs and 6-7 spiral cords, more often 6, Second whorl with 20 axial ribs and 7 spiral cords, third whorl with 22 axial ribs and 7 spiral cords, fourth whorl with 26 axial ribs and 7 spiral cords, fifth whorl with 32-33 axial ribs and 4 spiral cords. Body whorl with 32 axial ribs and 4 upper spiral cords. Suture strongly con- stricted, bordered by a strong, nodular, subsutiel spiral cord. Base imperforate, strongly conical and gently con- vex, ornamented by 15 nodular spiral cords, 5 of which form siphonal fasciole. Aperture elliptical, fusiform, nar- row at terminations. Peristome shiny, very thick and strongly expanded. Outer lip thick, with a broad posterior sinus, withl2 denticles, most anterior denticle more elongated. Parietal region strongly reflected, with inter- nal nodules. Columella gently concave, with two pro- nounced, rounded folds, the adapical fold slightly larger Siphonal canal short and narrow in distal extremity. Type Material: © Holotype, MZUSP 75932 [Length 22 mm, Width 8.3 mm]; juvenile shells: 1 paratype, ANSP 413550; 3 paratypes, MORG 50.658; 2 paratypes, MNR] LOTIS; 4 paratypes, MZUSP 78933. All from type local- 1S Noy. 2000. Type Locality: Northeastern Brazil, off the State of Pernambuco, 08°46.5' S, 34°44.5' muddy bottom, 690 im. Geographical Distribution: Continental slope off Pernambuco, 08°46.5' S, 34°44.5' W, northeastern Brazil. S. F. B. de Lima, J. C. N. de Barros and R. W. Petit, 2007 Page 101 Figures 1-5. 5 Gerdiella alvesi new species, holotype MZSP 78932, length 22 mm. 1. Apertural view; 2. Detail of aperture; 3. Lateral view showing profile of outer lip. 4. Protoconch. 5. View of ornamentation of second and third teleoconch whorls. Scale bars Figures 2, 3,5 = 500 pm; Figure 4 = 200 zm Etymology: Named in honor of Dr. Marcos Souto Alves of the Biology Department, in the field of Zoology at the Universidade Federal Rural de Pernambuco UFRPE) for having sent the first author under an in- ternship at the Malacology Laboratory of the UFRPE. Remarks: The protoconch of the holotype is not well illustrated as it is damaged. Nonetheless, we were able to characterize the protoconch of juvenile specimens of the new species (Figures 6-1] Gerdiella sp Figures 15-19 Material Examined: One damaged specimen MZUSP 78934. length 18.3 mm, off the state of Pernam- buco, northeastern Brazil, 08°46.5' S, 34°44.5' W, muddy bottom, 690 m, 18 Noy. 2000 Geographical Distribution: The Continental Slope off Pernambuco, northeastern Brazil. Remarks: The single specimen of Gerdiella sp. may represent a new species. However, we prefer not to name it as the shell is damaged and eroded. The proto- conch of this species has the same characteristics, and the same number of whorls, as Gerdiella alvesi. This specimen has two equal and very narrow columella folds, slightly different from those of G. alvesi. The first second and third post-nuclear whorls are rounded and ornamented by finely cancellated axial ribs and spiral cords, forming small nodules at their intersections, simi- lar to those of Gerdiella gerda and Gerdiella santa. The axial ribs are thicker than the spiral cords, (9 on the body whorl), as opposed to 6 for G. alvesi. The subsutural cord is weak on the first and second whorls of the teleoconcl Page 102 THE NAUTILUS, Vol. 121, No. 2 Figures 6-14. Juveniles of Gerdiella alvesi new species. 6-8. Protoconch under SEM, MORG 50.688; 9. SEM of protoconch of paratype MORG 50.688 showing microscopic spiral threads. 10-11. SEM of paratypes MORG 50.688. 12-13. Paratype, MZUSP 78933. 14. SEM of paratype MORG 50.688 showing growth lines. Scale bars: Figures 6—S = 200 wm; Figure 9 = 50 jm; Figures 10-13 = 500 jm; Figure 14 = 50 pm. The subsutural region is flat, encompassing the first and second spiral cords, which do not form a shoulder. There are seven spiral cords on the first and second whorls, and 10 on the third. There are 27, 30, and 36 axial ribs on the first, second and third post-nuclear whorls, respectively. In relation to G. alvesi, Gerdiella sp. has the same num- ber of spiral elements on the first and second post- nuclear whorls. This number is higher, however, on the third whorl. The number of axial ribs on the first three whorls of the teleoconch of G. alvesi is less than that of Gerdiella sp. DISCUSSION The conchological characters of Gerdiella sp. cannot be completely and conclusively compared to any of its con- geners until better material is collected for study at the species level. The analysis presented above distinguishes this species from the unnamed shell figured by Ver- hecken (2002: figs. 9, 10) by the greater number of axial and spiral ribs and threads on the first and second whorl. Gerdiella alvesi stands out from its congeners by the strong, uniform axial and spiral ornamentation, which forms large. rounded nodules that are quite pronounced pecially on the body whorl. The heavily thickened outer lip is very similar to that of Gerdiella gerda, prin- cipally on the sigmoid contour, and is lirated in the same way as Gerdiella cingulata, but not as coarsely as de- scribed by Olsson and Bayer (1972: 879). Two columellar folds are present on all the species. In both G. cingulata and G. alvesi the adapical fold is larger and there are no tubercles between the folds in the latter of the two spe- cies. In G, alvesi, there is no projection of the parietal callus on the outer lip, which is present in G. gerda and G. santa. Among the Gerdiella, the spire of G. alvesi has the least number of whorls and lacks varices. ACKNOWLEDGMENTS We are deeply grateful to: Enilson Cabral of the Re- search and Management Center of Fishing Resources of the Northeastern Coast—CEPENE/IBAMA—for his personal efforts in the collection of sediment from the Continental Slope off northeastern Brazil and for the donation of all this conchological material to the Mala- cology Laboratory of the Universidade Federal Rural de Pernambuco, Brazil; Fernanda Emanuele, Instituto Tec- noldgico de Pernambuco (ITEP) for taking the electron scanning micrographs; Mr. Richard Boike for an early translation of the text into English. Dr. M. G. Harase- S. F. B. de Lima, J. C. N. de Barros and R. W. Petit, 2007 Page 103 Figures 15-19. Gerdiella sp.. MZSP 78934, length = 18.3 mm. 15. Abapertural view. 16. Columellar folds under SEM. 17. View of ornamentation and subsutural flattening of the second and third teleoconch whorls. 18-19. Protoconch, Scale bars: Figures 16, 18, 19 = 200 pm; Figure 17 = 500 pm. wych, National Museum of Natural History, Smithsonian Institution, Washington, DC, kindly reviewed the manu- script. LITERATURE CITED Harasewych, M. G. and R. E. Petit. 1982. Notes on the mor- phology of Cancellaria reti: ulata (Gastropoda: Cancellari- idae). The Nautilus 96: 104-113 Harasewych M. G.. R. E. Petit, and A. Verhecken. 1992. Two new species of Cancellariidae (Gastropoda: Neogas- tropoda) from Brazil. The Nautilus 106: 43-49 Kaicher. $. D. 1978. Pack #19, Cancellariidae. Card catalogue of world-wide shells. Cards 1859-1964 Olsson, A. A. and F. M. Bayer. 1972. Gerdiella, a new genus of deep-water cancellariids. Bulletin of Marine Science 22 8$75-S80 Petit, R. E. and M. G Harasewych 1993. A new Mericella (Mollusca: Gastropoda: Cancellariidae) from northeastern Africa. Proceedings of the Biological Society of Washing- ton, LOG: 221-994 Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-Expedition II. Teil. Deutsche Tiefsee-Expedition 1598-1899, 17(2): 35-382, pls. 13-46. [Dual pagination; also numbered 1-348, pls. 1-34. | Thiele, J. 1929-35. Handbuch der systematischen Weich- tierkunde. Gustave Fischer, Jena. 2 vols. [1(1), 1-376 (1929): 1(2), i-vi, 377-778 (1931); 2(3), 779-1022 (1934) 9(4), i-vi, 1023-1154 (1935)]. Verhecken, A. 2002. Atlantic bathyal Cancellariidae (Neogas- tropoda: Cancellarioidea): Additional data and description of a new species. Journal of Conchology 37: 505-514 Verhecken, A. and L. Bozzetti. 2006. New data on East-African Mericella species, and description of a new species of Scalptia (Neogastropoda Cancellarioidea: Cancellariidae Gloria Maris 45: 14-25 THE NAUTILUS 121(2):104, 2007 Page 104 Book Review Land and Freshwater Molluscs of Brazil Luiz Ricardo L. Simone. 2006. Land and Freshwater Mol- luses of Brazil. Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo, Brazil. 390 pp., including over 1100 text- figures. ISBN 859066700-6. 8.5 by 11 in. Hardback; in En- ali sh. $125 from US booksellers: 92.5 to 94 € in Europe. This large-scale work is a timely and sorely needed rem- edy for one of the most stark deficiencies in the global inventory of molluscan biodiversity. The author has re- cruited no less than 70 identified collaborators and used the resources of 27 prominent institutional collections to formulate a presentation of 1074 native and 33 intro- duced species-level molluscan taxa inhabiting the land and fresh waters of Brazil and/or nearby areas. “The com- position of this native non-marine fauna (about 700 ter- restrial and over 370 aquatic, over 950 gastropods, and 116 pelecypods) invites comparison with other areas such as the USA, where freshwater clams are notably more speciose, and non-marine snails occur in roughly com- parable diversity. The work is organized thus: a one-page Introduction, four pages of legends and acknowledgements, a Table of Contents, which is a systematic outline of taxa treated, 278 pages of consistently formatted illustrations and companion text blocks in telegr. aphic style, a bibliography of 2696 titles, and an index of all taxa from phylum to species-level, the later presented with trivial name first. The format is simple and efficient. The Introduction indi- cates the method of presentation of taxa, each consecutively numbered in a conventional phylogenetic sequence (genera non- menage family unit, but species alphe ibetical within genus) The reiterative eye- -catching marginal icons (up to four) in each text block are a ad: geographic dis- tribution (blue globe); literature citations (re d printe sd page): source of companion figure, each of which is like-numbered (green eyeball); a non-critical synonymy (equal sign on or- ange); and “N.B.” for random notations when appende cl. The English diction is deficient on this page, but the author's intent is generally comprehensible. One defining feature of the work is enunciated in the In- troduction: Simone characterizes the figures as “normally based on type specimens.” Close perusal confirms this as the case, with name-bearing types from virtually all of the cited (27) institutional collections being depicted in dedicated pho- tographs. For the exceptions, it is apparent that paratypes, voucher specimens, iconotypes, and other levels of authen- icity were assiduously pursued and exploite od: for the slug groups this proved generally impossible. The photogr: iphs are generally of high fide lity, with appropriate magnification to acilitate identification. About three dozen photogr: iphic vi- gnettes, mostly of living snails, appropriately placed at various oints add a dime sion of vitality. Bibliographic citations are arr anged in near flawless alpha- vetical-chronological sequence and numbered consecutively. One or more such numbers appear in each text block, and an attempt is made to code them with one to six lower case code letters indicating if the work contained a description, figure, etc. Regrettably there are many omissions of the important “On code. which indicates “original,” as in description. The concermed reader must alphi ibetically search the bibliography using author and date expressed after the binomen at the heading of such entries. Perhaps unfortunately, the works of : ae and Pilsbry are particularly prone to this oversight. The fidelity of the citations appears to be excellent, although Bahiensis miliola (no. 591) ) appears to date from @’ Orbigny, 1837, rather than 1835 as stated (Pilsbry, 1901: 32: Sherborn and Griffin, 1934). A minor and easily remediable biblio- graphic nuisance is the lack of identification of the G. B. Sowerbys by generation, In the course of the work several generic reassignments (clearly marked as “n. comb.”) are install d and nomina nuda revealed. Species no. 1071, Byssanodonta riograndensis (Ihering and Morretes, 1949), is thus designated, but a pho- eee of an ANSP specimen accompanies this entry. Even though the Code eins 1999: Article 13.1.1), since 1930, has not recognized a binomen and figure indication in the ab- sence of a written description as basis for an available name— in this instance attributable to Simone—an explanation for his treatment of this apparent taxon would be welcome by the reader. Does Simone think this is a valid, unnamed taxon? The specimen figure on p. 309 captioned Lamellaxis clavinulus (Potiez and Michaud, 1838) ) appears not to be that species but L. micrus (@Orbigny, 1835), which is treated as Allopeas micra {sic] on p. 14. On p. 312, Europe is given as the origin of the non-native Bradybaena similaris (Férussac, 1821). It is more li kely from eastern Asia. There are technical problems with the typesetting such as wholesale deletion of dozens of single letters, particularly noteworthy on page 23, and the occasional misspelling or improper diacritical mark can be a minor distraction. The foregoing minor critique notwithstanding, Simone’s book is monumental. It is a prodigious work in both scope and the quality of the research. It is certain that all serious work- ers will find it indispensable in the understanding of the ex- tensive and complex Neotropical malacofauna. On another plane, its application to the analysis of other major world faunas will impel us to a better appreci ation of the systemat- ics, evolution, zoogeography, and macroecology of nonmarine Mollusca on a global scale. LITERATURE CITED International Commission on Zoological Nomenclature (ICZN). 1999. Intemational Code of Zoological Nomenclature. Fourth edition International Trust for Zoological Nomenclature, London, xxix + 306 pp Pilsbry, H. A. 1901-02. Manual of Conchology (second series). 14. Oriental bulimoid Helicinidae. Academy of Natural Sciences, Philadelphia. [iv] + 302 + xcix pp, 62 pls Sherborn, C. D. and F. J. Griffin, 1934. On dates of f pub lication of the natural history portions of Alcide d'Orbigny’s “Voyage dans Annals and Magazine of Natural His- tory series 13 1L0(13): 130-134. Harry G. Lee 1132 Ortega Forest Drive 32210 USA r Ame rique me sridioné ile Jacksonville, FI shells@hglee com | MBL WHO! Libr uni) 5 WH INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes articles on all aspects of the biology, ipaisentataue and systematics of mollusks. Manuscripts describing original, unpublished research and review articles will be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not require an abstract. 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Box 1580 Sanibel, FL 33957 THE@MNAUTILUS Volume 121, Number 3 October 10, 2007 CONTENTS ISSN 0028-1344 Gary W. Schmelz The Epitoniidae (Gastropoda: Ptenoglossa) from the lower Alum Bluff Roger W. Portell Group (lower to middle Miocene) of Florida, with descriptions of nine NEW SPSClES 5 ke alar aed SAGE OR NewS Se Mend Wc @ Aldean aiararin 105 Francisco M. Heralde III Molecular phylogeny of some Indo-Pacific genera in the subfamily Turrinae Maren Watkins H. Adams and A. Adams, 1853 (1838) (Gastropoda: Neogastropoda) ... . . 13] John-Paul Ownby Pradip K. Bandyopadhyay Ameurfina D. Santos Gisela P. Concepcion Baldomero M. Olivera Claudia Muniain Reproductive biology of the nudibranch Doris fontainei @Orbigny, 1835 Carlos S. Gallardo (Gastropoda: Opisthobranchia) from the Magellanic Region ........... 139 Pablo E. Penchaszadeh Juliana M. Harding Two modern records of the southern oyster drill Stramonita haemastoma M. G. Harasewych floridana in Chesapeake Bay, USA 2... 2. ee 146 Paolo Mariottini Brachycythara beatriceae, a new species from the Alboran Sea and the eastern Atlantic Ocean (Gastropoda: Neogastropoda: Conidae) ......... 159 Sponsored in part by the State of Florida, Department of State, Division of Cultural Affairs, the Florida Arts Council and the National Endowment for the Arts pRI> & . 9° 7, ~, Qe 78) ie) s ad o ZS a NATIONAL p x ENDOWMENT 0 ~ FOR THE ARTS THE NAUTILUS 121(3):105-130, 2007 Page 105 The Epitoniidae (Gastropoda: Ptenoglossa) from the lower Alum Bluff Group (lower to middle Miocene) of Florida, with descriptions of nine new species Gary W. Schmelz! 5575 Dogwood Way Naples, FL 34116 USA P.O. Box 117800 Gainesville, FI Roger W. Portell Florida Museum of Natural History University of Florida 32611-7800 USA portel@flmnh.ufl.edu ABSTRACT A comprehensive study of the family Epitoniid: ve that exists in the lower to middle Miocene portion of the Alum Bluff Group of Florida (USA) was conducted. A total of 14 species was examined. Of these, 12 are considered valid members of the family Epitoniidae. They include three previously described species, Amaea gardnerae, Cirsotrema cirritum, and Epitonium virginiae, plus nine new species. Seven of the new species were collected from the Chipola Formation and two from the Shoal River Formation. One new species is assigned to the genus Cirsotrema, two are assigned to the genus Opalia, and six are assigned to the genus Epitonium. Cirsotrema previously reported from the Chipola Formation as Cirsotrema dalli, a Pleistocene to Recent species, has been described as a new species. Epitonium alaquaense reported from the upper Miocene Choctawhatchee Formation (upper Alum Bluff Group) and questionably placed in the Shoal River Formation fauna by Gardner (1947) is no longer recognized as a Shoal River Formation species and Gegania acutissima has been placed with the Architectonica-like members of the family Mathildidae Additional Keywords: Miocene, Chipola Formation, Shoal River Formation, Cirsotrema, Opalia, Epitonium, Chipola River, Tenmile Creek, Farley Creek INTRODUCTION The family Epitoniidae has an extensive history with rep- resentatives dating back to the early Mesozoic. According to Clench and Tumer (1950), the group appears to have reached its peak of diversity during the Eocene and Mi- ocene epochs. In Florida (USA), members of the family are well represented in early Miocene deposits and, to a lesser extent, in middle Miocene deposits of the Alum Bluff Group. The Alum Bluff Group consists of five named strati- graphic units (Figure 1). From oldest to youngest these units are the Chipola Formation, Oak Grove Sond. Shoal ' Author for correspondence River, Choctawhatchee, and Jackson Bluff formations (Huddlestun, 1984). All of the Alum Bluff strata occur in outcrops in the western portion of the Florida panhandle (Figure 2). The lower Miocene Chipola Formation was depositec | approximately 1S mya (Jones et al., 1993) and outcrops along Tenmile, Farley, and Fourmile creeks, and the (¢ Chipola, Yellow, Choctawhatchee, and Apalachi- cola rivers. To date, most Chipola Formation specimens have been collected from Tenmile, Farley, and Fourmile creeks, and the Chipola River. Collections of fossil speci- mens from the Oak Grove Sand along the Yellow River and the Chipola Formation along the Apalachicola River are limited, given that these locations typically can only be accessed during very low water levels. According to Vokes (1989), Tenmile Creek, F arley Creek, and C hipola River complex alone encompasses over 7.5 miles (~12 kim) of Chipola Formation exposures. The middle Mi- ocene Shoal River Formation was de posite od approxi- mately 12 mya (Jones et al., 1993). All known Shoal River Formation outcrops are west of the Chipola Formation exposures with most of the collecting areas situated along the Shoal River in Walton County. Overviews of the ge- ology, stratigraphy, and paleontology of the Chipola aad Shoal River aa can be found, respectively, in Vokes (1989) and Portell et al. (2006). Gardner (1947), as part of her monographic treatment f the ae ee in fauna of the Alum Bluff Group, dis- cussed three species belonging to the family Epitoniidae. These species were Epitonium (Spiniscala) virginiae (Maury, 1910), Epitonium (Clathrus ) ) alaquae nse (Mans- field, 1935), and Gegania acutissima (Dall, 1892). How- ever, only two of these taxa are herein considered to belong to the family Epitoniidae. Epitonium virginiae was collect d from a Chipola Formation site along the east bank of the Aps ilachicola River. It was describe ad by Maury (1910) from a single, extremely small (3.7 mm maximum height x 1.5 mm maximum width) specimen that was part of the Cornell University Collection (now deposited at the Paleontological Research Institution). Epitonium alaquaense, recorded by Mansfield (1935) Page 106 THE NAUTILUS, Vol. 121, No. 1c) Cooke (1945) DUPLIN MARL (Ecphora and Cancellaria zones) Huddlestun (1984) JACKSON BLUFF FM (Ecphora and Cancellaria zones) Vokes (1989) JACKSON BLUFF FM (Ecphora and Cancellaria zones) Scott (2001) Carter et al. (2003) JACKSON BLUFF FM (Cancellaria zone) This Paper JACKSON BLUFF FM (Ecphora and Cancellaria zones) PLIOCENE CHOCTAWHATCHEE FM RED BAY FM Arca rubisiniana zone (Arca zone) (Arca zone) CHOCTAWHATCHEE RED BAY FM —M A (Arca zone) (Arca zone) Upper (Yoldia zone) Yoldia waltonensis = ir or w 2 x = = 9 Ed w Middle (Main shell bed) MIOCENE SHOAL RIVER FM Glycymeris waltonensis zone WHITES CREEK MEMBER YELLOW RIVER FM (Yoldia zone) SHOAL RIVER FM YELLOW RIVER FM CHOCTAWHATCHEE FM. (Yoldia zone) SHOAL RIVER FM (Includes “Cardium beds” of Gardner (1926), Yoldia and Glycymeris zones, and Whites Creek Member) ALUM BLUFF GROUP SHOAL RIVER FM ALUM BLUFF GROUP (UNDIFFERENTIATED) WHITES CREEK BEDS Cardium taphrium zone OAK GROVE SAND CHIPOLA FM CHIPOLA FM Figure 1 from the Arca zone of the Choctawhatchee Formation, is late Miocene (Huddlestun, 1984). When Gardner (1947) collected a shell fragment similar to Mansfield’s shell at a Shoal River site she included it as part of the lower Alum Bluff Group as defined by Cooke (1945). The col- lective evidence now suggests that E. alaquaense should Zz not be listed as part of the fauna of the lower Alum Bluff Group (see Figure 1). The reasons for this are twofold. First, an extensive examination of the Chipola and Shoal River formation fossils in the Florida Museum of Natural History (including Tulane University and Florida Geo- logical Survey enlle sctions), Paleontologic: il] Research In- stitution, Smithsonian’s National Museum of Natural History, Museum of Natural History, The Academy of Natural Sciences, Alabama Ge ologic: ul Sur- vey, and private collections during, this study did not American uncover a single specimen that could be referred to as E (1947: from * no7 577) stated that she shell fragment ‘a horizon. slightly than the Based upon this remark and that no additional specimens simi- lar to E mentioned collections, it seems fair to assume that Gard- alaquae nse. Second, Gardner collected her higher typical Shoal River formation.” alaquaense were ever found in the above- ner s specimen be longe d to strati tyounger than the Shoal River Formation; most probably to the upper Miocene Choctawhatchee Formation Dall (1892) described Tuba acutissima and placed it in the family Mathildidae Tuba Gardner (1947) replaced the ge- nu with Gegania and tentatively assigned it to the a OAK GROVE FM. /, ae CHIPOLA FM Alum Bluff Group stratigraphic nomenclatural history and correlation (in part). x OAK GROVE SAND OAK GROVE SAND CHIPOLA FM CHIPOLA FM Modified from Portell et al. (2006), family Epitoniidae because she felt that its morphologi- cal characteristics more close ly coincided with the sub- order Ptenoglossa. Gardner's generic assignment of Ge- gania was based upon the shell similarity to Gegania pinquis Jeffreys, 1884, a species collecte d during the Porcupine Expedition off Cape Mondego, Portugal. However, the genus Gegania has since been assigned to Architectonica-like members of the family Mathildidae (Vaught, 1989). In addition to the three species of Epitoniidae listed for the Alum Bluff Group, Gardner (1947: 575) reported shell fragments belonging to “at least a dozen” Gardner stated that most of the specimens were so im- species. pe rfectly preserve od that only subge meric determinations could be made. Four of the unidentified « :pitoniids came from the Aldrich Collection (housed at Johns Hopkins University) and the remaining species were from Gard- ner’s collection, Eleven of the fragments were obtained from Shoal River Formation localities and two were col- lected from Chipola Formation sites. Gardner placed eleven of the fragments in the genus Epitonium, one in the genus Scalina, and one in the genus Gegania. Six were placed in the subgenus Hyaloscala, two in Spinis- cala, three in Cinctiscala, and one in Nodiscala. [t is apparent from Gardner's (1947) discussion on the Epitoniidae of the Alum Bluff Group that future work remained to be done on the family. ae her publica- tion, three additional species of EF pitoniid ie have been reported from the Chipola Formation. These are Cir- G. W. Schmelz and R. W. Portell, 2007 Page 107 EE Jackson Alum Bluff Group Figure 2... Map of Florida showing counties with Alum Bluff Group sediments found at or near the surface. sotrema dalli Rehder, 1945; Scalina gardnerae (Olsson, 1967): and Cirsotrema cirritum Duerr, 2004. Cirsotrema dalli is an extant species reported to have undergone little morphological change since the early Miocene (Ols- son, 1967). Scalina gardnerae was described by Olsson (1967) from a single large specimen collected from Mc- Clellan’s Farm in Calhoun County, Florida. It is a fairly common species, and since its discovery, it has been found at numerous Chipola Formation sites by the au- thors and other investigators. Cirsotrema cirritum is a much less common species collected at a few sites along Tenmile and Farley creeks and the Chipola River (Du- err, 2004). Scattered among the material in museums and private collections are a large number of epitoniids from the Chipola and Shoal River formations of Florida that have been amassed over the last fifty years. Since many of the recently collected specimens are in excellent shape, we have undertaken the task of identifying and describing the new species and providing better descriptions and updated taxonomic placements of existing ones. As will be evidenced in this paper, many of these epitoniids have shell structures that are similar to those of extant and fossil species from Florida, the Caribbean, westem Eu- rope. and the eastern Pacific. According to Clench and Tumer (1951), DuShane 1979). Kilburn (1985), Nakayama (2003), Robertson (1983a; 1983b; 1993), Weil et al. (1999), and others, cur- rent classification of the Epitoniidae is based upon shell characteristics. For this study, morphological features such as shell size (height and width), number and shape of the nuclear whorls, number and shape of teleoconch whorls, number and placement of the costae on the body whorl, presence or absence of varices, type of sculpturing on the body whorls, shape of the aperture, shape and thickness of the outer lip, depth of the suture, presence of punctae, and the spire angle were used to help differ- entiate species. Spire angles were measured from pho- tographs. A vertical line was drawn through the axis of the shell and a protractor was used to determine the angle between the margins of the body whorls on both sides of the shell. Additionally, in order to ascertain whether or not the Chipola Formation species of Cirsotrema is the same as the Recent Cirsotrema dalli, we closely examined the varices on both forms. As noted by Weil et al. (1999) and others, the presence or absence of varices can be a key diagnostic feature for some species of Epitonium. We believe that the structural configuration and the number of varices that appear within a specific generic group, such as in certain Epitonium, will change over geologic time and that this morphological difference between the Recent and fossil forms is significant enough to justify naming the fossil form as a separate species. In part, this Page 108 THE NAUTILUS, Vol. 121, No. 3 decision to use the changes observed in varix count as a diagnostic feature at the species level is based upon the precedence of using the number, shape, and configura- tion of varices as a method of identifying different : spe- cies and genera in the family Muricidae. This technique has been used broadly by muricid gastropod investigators (Herbert, 2005, and references therein). Besides examining the varices on Cirsotrema dalli, we also abraded the external portion of an uncatalogued Re- cent shell in order to compare its underlying sculpture with that of its fossil counterpart. Even though many of the epitoniids examined are in excellent shape, the task of classifying them and placing them into their appropriate generic and subgeneric groupings was challenging. This is, in part, because there is very little natural history information on Recent spe- cies that provide insight into how these animals grow and how their growth is ee by environmental condi- tions. Subsequently, we have relied heavily upon the combined works of numerous past investigators to help us with this decision making process. These researchers included, but were not limited to: Brunet (1995), de Boury (1909), Clench and Turner (1950; 1951; 1952), DuShane (1979: 1988), Gardner (1947), Kilburn (1985), laa (2003), Robertson (1983a; 1983b; 1993), Weil et al. (1999), and Woodring (1959). At the generic and subgeneric levels of classification many investigators have y widely div ergent opinions about in which group a specific epitoniid should be placed. According to Clench and Turner (1950), de Boury's work on E pitoniidae lk left behind a long list of generic and subgeneric names with only the types avail: ble for diag- nostic analysis. This list includes seven generic and 19 subgeneric names (Weil et al., 1999). It was Clench and Turner's (1950) opinion that de Boury became confused about the overlapping characteristics of the Epitoniidae and rather than trying to place them into existing catego- ries, he established new genera and subgenera for them. To date, this confusion with ove lapping characteristics appears to have continued with the list of generic and subgeneric extant Epitoniid ae alone being expanded to 34: and; 35 names, respective ly (Weil et al., 1999). For our classification purposes we have decided to adhere, as closely as possible, to the more conservative phylogenetic scheme followed by Clench and Turner (1950) rather than the more rece nntly expanded version used by Weil et al. (1999) and Nakayama (2004). With fossils, placement of certain epitoniids into ap- propriate generic and subgeneric groups has been ham- pe re va by iC rosion ( { taphonomic degrad: ition). This process removes surface sculpturing present on living specimens and exposes subsurface characteristics that are remark- ably different. In this paper, an example of the impact erosion has on the external features of a shell is ilhas- trated with the new species Epitonium conwaiae. We have tried to survey all the pertinent descriptions and illustrations of both fossil and Recent « »pitoniids. For taxonomic Comparison purposes the most significant lit- erature came from publications dealing with the descrip- tions of Recent and fossil species collected from the United States, Central and South America, the northwest Atlantic, European continent, and the eastern Pacific. Institutional abbreviations used herein are: USNM: National Museum of Natural History, Smithsonian Insti- tution, Washington, DC; UF: Florida Museum of Natu- ral History (FLMNH), University of Florida, Gainesville; TU: Tulane University, (formerly housed in New Or- leans, Louisiana and now housed at the FLMNH):; PRI: Paleontological Research Institution, Ithaca, New York; ANSP: T The Academy of Natural Sciences, Philadelphia, Pennsylvania; and BMSM: The Bailey-Matthews Shell Museum, Sanibel Island, Florida. Because of privacy rights of landowners, specific locality data for specimens described below are not given. However, specific locality information is available to qualified researchers upon written request to portell@flmnh.ufl.edu. SYSTEMATICS Superfamily Janthinoidea Lamarck, 1812 Family Epitoniidae Berry, 1910 Genus Amaea H. and A. Adams, 1853 Type Species: by subsequent designation, Scalaria magnifica Sowerby, 1544. Subgenus Scalina Conrad, 1865 Amaea gardnerae (Olsson, 1967) (Figures 3-9) Description: Shell large, turriculate; protoconch miss- ing; nine or ten teleoconch whorls. Spire angle 19.5°, W hors convex, relatively thin, with cancellate sculpture. Cancellate pattern created by four spiral cords crossed by smaller, evenly spaced axial costae: square spaces within cancellate sculpture possess textured pattern cre- ated by fine overlapping axial and spiral threads. Suture deeply impressed. B Basal cord well-defined, surface slightly elevated and sculptured with thin spiral and axial hreails: Sculpture below basal disc lacks elevated spiral cords. Umbilicus absent. Holotype: USNM 645180, maximum height 47.5 mm, maximum width 14.4 mm. Type Locality: McClellan Farm, Calhoun County, Florida. Other Material Examined: UF 117045, | shell, Ten- mile Creek 04 (CA020) (= Tulane University locality TU 51), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation; UF 117087, 1 shell, locality and formation same as pre- ceding; UF 91459, 16 shells, Tenmile Creek 03 (CAQOI7) = Tulane University locality TU 546), Clarksville Quad- rangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation. Distribution: Amaca gardnerae is a fairly common Chipola Formation species. [t is abundant at several col- G. W. Schmelz and R. W. Portell, 2007 Page 109 Figures 3-9. Amaea gardnerae (Olsson, 1967). 3-4. Apertural and abapertural views of holotype USNM 645150 originally named Scalina gardnerae Olsson, 1967. Since its original description the specimen has degraded; maximum height 47.5 mm, maximum width 14.4 mm. 5-8. Apertural, lateral, abapertural, and basal views of UF 117045 shown for comparison to holotype (USNM 645150) and other Chipola and Shoal River formation epitoniid species; maximum height 17.55 mm, maximum width 5.4 mm. 9. Magnified view of sculpture of teleoconch of UF 117045. Scale bar = 0.6 mm lection sites along Tenmile Creek but is less abundant along Farley Creek and the Chipola River. Thus far, there are no reports of it having been collected along the Yellow or the Apalachicola rivers Etymology: Named for Julia A. Gardner, a pioneer researcher on Chipola Formation mollusks Discussion: Gardner (1945) reported an epitoniid fragment from the Chipola Formation which she as- signed to the genus Scalina. Olsson (1967) later de- scribed this species as Scalina gardnerae. A more thor- ough analysis of additional specimens of S. gardnerae now indicates that it should be placed in the genus Amaea. DuShane (1988) noted that members of the ge- nus Amadea are larger than any k nown Scalina and have a less distinct basal cord. In addition, DuShane observed that in the genus Amaea, the sculpture above and below the basal cord is different. Unfortunately, the poor con- dition of the type specimen described by Olsson makes it difficult to tell what the sculpturing was like in the vi- cinity of the basal cord Subsequent specnhnens, however have revealed that the sculpturing above and below the basal cord is different in S. gardnerae (Figures 5, 8) Clench and Tumer (1950), Weil et al. (1999), and Nakayama (2003) have all placed epitoniids with the shell sculpture described by Olsson (1967) for Scalina gard- nerae into the genus Amaea. We are in complete agree- ment with this placement and have assigned Olsson’s epitoniid to that genus Weil et al. (1999) has identified eight subgeneric forms of the genus Amaea. These subgenera are distinguished from one another by the types ol sculpture that appeal above and below the basal cord. Nakayama (2003) in hi: review of northwest Pacific epitoniids retained six of th« subgenera listed by Weil et al. (1999). Among thi subgenera listed for the genus Amaea by Nakay 2003), the present authors have assigned the Chipola ’ ] Formation species to the subgenus Scalina. According to Page 110 THE NAUTILUS, Vol. 121, No. 3 Nakayama (2003) and Weil et al. (1999) members of this subgenus possess convex body whorls with a cancellate sculpture of spiral cords and axial ribs. Representatives of the genus Amaea have been re- ported from a number of other fossil locations. These include Scala (Opalia) reticulata Martin, 1904, from the Miocene Calvert Formation of Maryland, Amaea (Scalina) ferminiana (Dall, 1908) from the Pliocene Es- meraldas beds of Ecuador, Scala (Acrilla) wiegandi (Bose, 1910) from Mexico and the Miocene Chagres Formation of Panama (formerly Canal Zone), Scalina pseudolerogi (Maury, 1925) from the Pliocene of Trin- idad, Epitonium (Ferminoscala) manabianum (Pilsbry and Olsson, 1941), and Epitonium (Ferminoscala) ele uth- erium (Pilsbry and Olsson, 1941) from the Pliocene fauna of western Ecuador, Scalina boylae (Olsson, 1967) from the Pliocene Tamiami Formation of Florida, and Scalina kendacensis Jung, 1971, from the Miocene Ken- deace Formation of Carriacou. Amaea ferminiana, origi- nally described from the Recent of Baja California, is an offshore species. It ranges from Mexico south to Peru (Weil et al., 1999). DuShane (1988) considered E. eleutherium and S. wiegandi to be synonymous with A. ferminiana and suggested that S. pseudolerogi as well as some other fossil species of Amaea may be synonymous with A. ferminiana. Comparison of A. gardnerae with A ferminiana clearly illustrates that the two species are et synonymous. Amacea fe rminiana possesses six to nine spi- ral cords of uniform thickness on the body whorls, while A. gardnerae possesses four broad primary cords with fae cords in between. The costae on A. gardnerae are also less prominent then they are on A. ferminiana. Com- parison of Amaea mitchelli (Dall, 1896), a Recent west- ern Atlantic species, to A. gardnerae was also made. Amaea mitchelli has 6-7 primary spiral cords on the body whorls (four of which are closely spaced below the whorl mid-line and two to three that are evenly spaced above) as compared to A. gardnerae which has four evenly spaced primary spiral cords. Genus Cirsotrema Morch, 1852 Type Species: monotypy. Scalaria varicosa Lamarck, 1822, by Cirsotrema chipolanum new species (Figures 10-14) Description: Shell large, turriculate, protoconch miss- ing; seven to eight teleoconch whorls. Spire angle 27°. Whorls slightly convex, strongly shouldered and joined. Primary unde rlying sculpture present on te sleoconch whorls consists of toy elevated vertical ribs. Suture deep, covered by exte “mal eal Seventeen to 23 sinuous, inclined, feather-like, broad costae on body whorls. Edges of foliated costae occasionally touch the preceding ones giving the shell surface a pitted appear- ance. In other instances foliated costae are slightly sepa- rated from one another. When feather-like costae sepa- rate from each other, five slightly elevated spiral cords seen On body whorls. Spiral cords, and Spaces between them, possess numerous distinct spiral threads. Thin ver- tical threads intersect spiral threads, creating faint can- cellate pattern. Varices formed from accretion of foliated costae; varices poorly developed and only slightly el- evated. Apical end of costae with pointed nodes. Surface of costae pitted with obliquely arranged small holes. Three or four obliquely arranged, slightly elevated, nar- row cords traverse each costa. Oblique cords on costae less pitted than remainder of surface area. Base of each costa stem-like, attached to a broad, crenulated, basal cord. Basal disk composite, created by a large outer basal cord with crenulated edges; a middle circle of narrow linear pits; and an inner columellar cord with a crenu- lated edge. Aperture subcircular. Columella short and arched. Lip margin thickened, pitted in unworn speci- mens. Less anode d specimens possess a slightly elevated node on the posterior outer margin of the ‘lip. Holotype: UF 117088, maximum height 32.9 mm, maximum width 15.0 mm. Type Locality: Tenmile Creek 03 (CA017) (= Tulane University locality TU 546), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation, Paratypes: UF 117089, | shell, locality and formation same as holotype; UF 91490, 5 shells, locality and for- mation same as ee UF 84575, 7 shells, Tenmile Creek 01 (CA002) (= Tulane University locality TU 830), Clarksville eee USGS 7.5’ Series (1945), Cal- houn County, Florida, Chipola For mation; UF 95161, 1 shell, Tenmile Creek 04 (CA020) (= Tulane University locality TU 951), Clarksville Quadrangle USGS 7.5’ Se- ries (1945), Calhoun County, Florida, Chipola For ma- tion: UF 85383, 1 shell, Chipola 09 (CAOIS) (= Tulane University locality TU 547), Clarksville Quadrangle USGS 7.5’ Series (1945), C Cahous County, Florida, Chipola Formation; UF 13966, 1 shell, Chipola 03 (CA005), Chipola River (exact collection site unknown), Calhoun County, Florida, Chipola eye UF 94650, 1 shell, Tenmile Creek 02 (CA003) (= Tulane University locality TU 70), Altha West Omdniile USGS 7.5’ Series (1982), Calhoun County, Florida, Chipola ares UF 84444, 1 shell, Farley on 03 (CA009) (= Tulane Unive rsity locality TU 825), Clarks- ville See USGS 7.5’ Series (1945), Gaon County, Florida, C bape “eae om UF 91356, 1 shell, Chipola O07 (CAOI5) (= Tulane University locality TU 554), Clarksville Guadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation; USNM 534489, 1 shell, Tenmile Creek 01 (CA002) (= Tulane University locality TU $30), Clarksville Quadrangle USGS 7.5' Series (1945), Calhoun County, Florida, Chipola Formation; USNM 534490, 2 fragments, locality sail formation same as pre ceding; U 5 M 534491, 2 frag- ments, Tenmile Creek 05 (CA021) (= Tulane Unive rsity locality TU 998), Clarksville Quadr ee USGS 7.5! Se- ries (1945), ( ‘lhieniyi County, F lorida, G hipola Forma- tion: USNM 534492. 1 shell, F arley Creek 03 (CA009) G. W. Schmelz and R. W. Portell, 2007 Figures 10-19. Cirosotrema chipolanum new species and Cirsotrema dalli Rehder, 1945. 10-13. Cirsotrema chipolanum a tural, lateral abapertural and basal views of holotype UF 117088; maximum height 32.9 mm pel i maximum width 15.0 mm. 14. Magnified view of s¢ ulpture of teleoconch of holotype UF 117088. Scale bar = 2.85 mm. 15-18. Cirsotrema dalli Rehder, 1945 apertural, lateral. abapertural, and basal views of UF 238698; maximum height 32.8 mm and maximum width 13.8 mm. 19. Magnified view of sculpture of teleoconch of UF 238698. Scale bar = 5.0 mm. UF 238698 live collected off Egmont Key, Pinellas Count Florida at about 52 m depth o Page 112 THE NAUTILUS, Vol. 121, No. 3 = Tulane Univ ersity locality TU 825), Clarksville Quad- eee USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation; USNM 534493, 10 shells, Tenmile Creek 03 (CAO17) (= Tulane University locality TU 546 and USGS 2212 “one mile west of Bailey's Ferry”), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhousi County, Florida, Chipola ep teen USNM 534494, 8 shells, same locality and formation as preced- ing, Distribution: Cirsotrema chipolanum is a fairly com- mon species. It is locally abundant at several Chipola Formation collecting sites along the Chipola River and Tenmile and Farley creeks. The fact that it has not been reported from the Oak Grove Sand along the Yellow River or from Chipola Formation sites along the Apalachicola River may simply reflect the degree of dif- ficulty collecting these localities at the appropriate peri- ods of low water level. Etymology: Named for the Chipola River. Discussion: The genus Cirsotrema has an extensive evolutionary history. Sohl (1964) established the genus Striaticostatum to represent a Cretaceous form of Cir- sotrema that lacked the faint spiral striations on the body whorl. By the Eocene the genus Cirsotrema was well established with numerous species being described from different localities (Dockery, 1980; Harris and Palmer, 1946; Palmer, 1937). C urrently, among extant forms, two species of Cirsotrema exist in ‘the western Atlantic (Weil et al., 1999). These are Cirsotrema dalli Rehder, 1945 and Cirsotrema pilsbryi McGinty, 1940. Two species of Cirsotrema have also been reported from lower Miocene Chipola Formation. Olsson (1967) identified a specimen collected along the west bank of the Chipola River south of Tenmile Creek as C. dalli and more recently Duerr (2004) described Cirsotrema cirri- tum from material collected at several different Chipola Formation sites. Olsson (1967) stated that after close examination of the Chipola Formation Cirsotrema he found practically no differences between it and the ex- tant species currently living in the western Atlantic. Since Olsson’s (1967) analy sis of the ¢ Chipola Cirsotrema, over 50 additional specimens have been collected from a va- riety of Chipola locations. When most of these specimens were closely scrutinized it became apparent to the present investigators that the Chipola Formation Cir sotrema studie d by Olsson (1967) was not C. dalli. As noted by Clench and Turner (1950), the sculptur- ing on the body whorls of representé oe. of the genus Cirsotrema, is extremely complicated. It was suggested by Clench and Turner, and observed in the present Pure that two layers of sculpturing are prese nt on the body whorls of Cirsotrema dalli. There is an outermost one that consists of foliated costae that may or may not join each other along their convoluted e dee »s and asec ‘ond: uy layer of aeinare cd costae and spiral cords. Because of this dual type of sculpture Clench and Tumer warned against identifying different species of Cirsotrema from beach worn specimens. Often with fossil specimens it is difficult to find shells that have not been eroded. However, a sufficient number of well-preserved Chipola Formation Cirsotrema were discovered which allowed a more detailed comparison between the fossil form of this genus and its Recent counterpart. These detailed studies lead to the following observations: First, when the underlying sculpture of the Chipola Formation Cirsotrema was exposed no difference was discovered between it and the underlying sculpture ex- hibited by C. dalli. Second, although varices are present on both the Chipola Formation Cirsotrema and its Recent counter- part, the varices on the Chipola Formation species are clearly not as well developed as those of Recent C. dalli. In fact, the varices on C. chipolanum are at times so poorly formed that it is difficult to identify them as va- rices. In C. dalli the costae are fused together to forma pronounced, elevated ridge (Figures 15-19) that, in well- preserved specimens, ee a slightly crenulated margin. Conversely, with C. C. chipolanum the varices consist of little more than one or two slightly raised costae (see Figures 10-13). In addition, even in instances where two costae are fused together to form a varix, the fusion is often not complete and a distinct line of demarcation can be seen. Third, when a comparison of the number of varices in relation to the height of the shell was undertaken with well-preserved specimens of C. dalli and C. chipolanum, and a least squares regression analysis was done on the two species, the qeaulis showed a significant correlation between the height of the shell sel the number of va- rices with C. dalli (0.922), and a low correlation between shell height and the number of varices with C. chipo- lanum (0. 499). Table 1 below provides the statistical re- sults of this study and a comparison of the number, size range, average size, varix range, and average number of varices of the specimens | in the analysis. As a result of the 1 regression analy sis tw o other differ- ences between the varices of the two species was also noted. Although the varices appeared at random intervals on the body + of both species, in C. chipolanum the first varix did not appear until after the third teleoconch whorl, while in C. dalli varices would appear just after the protoconch. In addition, when the number of varices Table 1. Results of least squares regression analysis compar- ing shell he sight with the number of varices found in C. dalli and C. chipolanum. C. dalli C. chipolanum No. of specimens examined 23 16 Correlation coefficient 0.922 0.499 Range of shell height (mm) 5.8-37.5 5.0-44.2 Mean height (mm) 15:31 17.2] Range of varices 2-27 0-10 Mean no. of varices §.82 4.75 G. W. Schmelz and R. W. Portell, 2007 Page 113 was compared between the different species it was discov- ered that C. dalli, on average, had significantly more varices than C. chipolanum [X = 8.82 vs. 4.75]. Table 2 below summarizes the structural differences in shell morphology between C. dalli and C. chipolanum. In addition to comparing the differences between the shape and number of varices between species, the table also compares the number of teleoconch whorls, the number and arrangement of costae, the spire angles, the arrangement of the costae, and the placement of varices on the body whorls. Another point worth noting is that C. dalli and C. chipolanum came from very different environments. Cir- sotrema dalli lives in cooler continental shelf waters at depths of 37 to 227 m (Clench and Turner, 1950) while C. chipolanum thrived in a shallow, warm, tropical, reef habitat. Although not a great deal is known about how the environment and food supply impact the growth of wentletraps [Robertson (1953a) and Weil et fal (1999)], there has been sufficient studies done on the family Mu- ricidae (Vokes, 1973) that will allow us to speculate as to the reasons for the number and structural differences observed between the varices of C. dalli and C. chipo- lanum. According to Vokes (1973) varix (axial growth ridge) development in muricids occurs during resting stages in shell formation. These stoppages may come about as the result of food shortages or perhaps as the result of environmental changes. The weaker develop- ment and lower number of varices in C. chipolanium may indicate that this species lived in a rather stable environ- ment where there were a large eee of prey species for it to consume. Conversely, dalli has more and better developed varices ou: stoppages) which may be due to lack of prey or perhaps increased predation pressures. In all probability, C. chipolanum is the ancestral form of C. dalli. It is the opinion of the investigators that C. chipolanum probably retreated from the coastal waters ste the deeper offshore waters during intervening ice ages where it gradually evolved into the extant, and mor- phologically ciendlae: C. dalli. Clench and Turner (1950) did not assign a subgeneric name to this genus even though de Boury (1909) did. Table 2. A comparison of the structural differences in shell morphology between C. dalli and C. chipolanum. dalli C. chipolanum No. of specimens examined 23 16 Teleoconch whorls 9-10 7-8 Spire angle 26.5° 27.0° Costae arrangement touching sometimes separated Mean no. of costae 20.6 19.] Range in costae number 18-23 17-2] Range in varix number 2-27 0-10 Mean varix number 8.82 4.75 ard start on 3 body whorl start on 1** body whorl Varix placement Clench and Turner's reason for not using subgenera is not stated, but in all probability the authors did not think that structural differences between members of this ge- nus were sufficient to warrant their use. Since Clench and Turner's publication, Weil et al. (1999) and Nakayama (2003) have assigned a number of subgenera to the genus Cirsotrema. It is interesting to note that between these authors there is not complete agreement as to which subgenera are valid, For example, ene oon aoa the pees Boreosc sea to de- Hie: like the W He W eil et al. ( 1999) 1 raised fe sub- genus Boreoscala to the generic level. In addition, Weil et al. questions the use of Elegantiscala as a subgenus of Cirsotrema while Nakayama retained this subgenus and used it for many of the species of Cirsotrema he de- scribed from the northwest Pacific. The present authors have followed the more conser- vative gee used by Clench and Turner (1950), Ab- bott (1974), Duerr (2004), and Landau et al. (2006) and have not nome a subgenus to the new Chipola For- mation species. Cirsotrema togatum (Hertlein and Strong, 1951) i morphologically sionilay to C. chipolanum. Duerr (5004) considers C. togatum to be a westem cognate of C. dalli. Cirsotrema togatum has also been reported from the Pliocene Esmeraldas beds of northwestern Ecuador by Pitt (1981) and DuShane (1988). Cirsotrema chipolanum differs from C. togatum in that it has fewer and less distinct spiral cords on the body whorls and the varices are much less developed. Cirsotrema woodringi Olsson, 1967, reported from the Miocene Gatun Formation of Panama and the Pliocene Tamiami Formation at Sunnyland, Florida, is similar to C. chipolanum. The main difference between C. woo- dringi and C. chipolanum is that C. woodringi lacks the inclined spiral sculpture that is present on the flat outer surface of the costae in C. chipolanum. Cirsotrema cirritum Duerr, 2004 (Figures 20-28) Description: Shell medium to large. Turriculate. Much of protoconch missing. Last protoconch whorl transitions from a smooth surface into wavy axial lamellae that gradually enlarge into thickened foliated costae. Spire angle 97° Eight strongly shouldered teleoconch whorls present. Suture deep. Thirteen crenulated, re- curved, axial costae present on last body whorl. Apical ends of axial costae sharp. Costae made up of multiple wavy lamella that possesses a fine irregular diamond- shape ‘d pattern that is inclined adaperturally. Costae separated by wide intercostal spaces. Intercostal spaces possess five ‘rounded spiral cords. Cords and intervening spaces have a cancellate sculpture created by overlapping vertical and horizontal threads. Anterior reflected pro- jections of costae on last body whorl form a basal ridge with a crenulate outer margin. Costae on basal ridge not fused. Varices absent. Columella short and arched. Ap- erture subcircular. No umbilicus. Page 114 THE NAUTILUS, Vol. 121, No. 3 Figures 20-28. Cirsofrema cirritum Duerr, 2004. 20-23. Apertural, lateral, abapertural, and basal views of holotype UF 110972; Jor mum height 28.5 mm, maximum width 11.8 mm, 24. Magnified view of sculpture of teleoconch of UF 110972. Scale bar = 2.75 9 5-28. Apertural, lateral, abapertural, and basal views of paratype UF 67746; maximum height 29.6 mm, maximum width 13.4 Note: Arrows point to apparent varices on paratype UF 67746 thus based upon description by Duerr (2004, p-. 154-155) a key diagnostic feature of no varices, this paratype was mistakenly identified. In ow opinion UF 67746 is representative /,; ] Fa Upolanun Tre species G. W. Schmelz and R. W. Portell, 2007 Page 115 Holotype: UF 110972, maximum height 28.5 mm, maximum width 11.8 mm. Type Locality: —Tenmile Creek 01 (CA002) (= Tulane University locality TU 830), Clarksville os eae USGS 7.5’ Series (1945), Calhoun County, Floves, Chipola Formation. Note: Duerr (2004) erroneously listed the GPS coordinates of 29° 30.05’ N, 85° 11.00’ W for this locality and provided no datum. These coordi- nates are approximately 17 km south of Saint Vincent Island, Florida in the Gulf of Mexico. Additionally, Du- err (2004) listed the type locality as in the SE1/4 of Sec. 7. It should have read SEI/4 of Sec. 12. Other Material Examined: UF 112019, paratype, Farley Creek 03 (CA009) (= Tulane University locality TU 825), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation; UF 67746, paratype, locality and formation same as pre- ceding; BMSM 15301, paratype, Tulane University a cality TU 458, Chipola pariew USNM 534499, shells, Tenmile Creek 03 (CA017) (= Tulane Univ ersity locality TU 546), C lanksville Deere USGS 7.5’ Se- ries (1945), Calhoun County, Florida, Chipola oe tion. Etymology: The name cirritum is derived from the Latin cirrus meaning “filamentous” which refers to the fringed costae of this species. Discussion: One of the key diagnostic features of C. cirritum is its lack of varices. However, when comparing the paratypes of C. cirritum with the holotype, one paratype, identified as C. cirritum, was discovered to possess varices that were similar to those observed in C. chipolanum (see Figures 25-28). This discovery gener- ated some confusion and resulted in an exhaustive inves- tigation to determine if only one species of Cirsotrema (C. cirritum) existed in the Chipola Formation or if there were two distinct species and a mistake had been made with the selection of one of the paratypes of C. cirritim. The conclusion drawn from this analysis was that there are two different species of Cirsotrema in the Chipola Formation and that the C. cirritum paratype (UF 67746) was mistakenly selected. When examining the different species of the Chipola Formation Cirsotrema it is easy to see how this mistake occurred. The varices on C. chipolanum sometimes can be easily overlooked without careful examination under a microscope. A summary of differences between C. cir- ritum and C. chipolanum is found in Table 3 Table 3. A comparison of the structural differences in shell morphology between C. cirritum and C. chipolanum. C. cirritum C. chipolanum Mean varix number 0 4.75 Costae number 13 17-21 Foliated costae widely separated closely packed Basal ridge costae not fused costae fused Cirsotrema cirritum is a fairly rare species that has been obtained from only three fossil localities: one each on the Chipola River (TU locality 458), Tenmile Creek (TU locality 951), and Farley Creek (TU locality $25 As Duerr (2004) noted, C. cirritim is a fairly a e tive species that bears some resemblance to several types of Recent Indo-Pacific Epitoniidae. Among the compa- rable Recent species, Duerr (2004) reported were Cir sotrema plexis Dall, 1925, Cirsotrema fimbriatulum (Masahito et al., 1971), Cirsotrema rugosum (Kuroda and Ito, 1961), and Cirsotrema excelsum Garcia, 2003. Among fossil species C. cirritum is most similar to the Miocene species Cirsotrema undulatum (Jung, 1965) from the Paraguana Peninsula, Venezuela. Cirsotrema undulatum is a medium-size shell with six to eight post- nuclear whorls that possess thin, widely sp: aced axials (Jung, 1965). Cirsotrema cirritum differs from C. undu- latum by having fewer axial costae on the body whorls (13 versus 17-21) rer five spiral cords on each ‘body whorl versus four. The presence of two species of Cirsotrema in the tropical Chipola Formation environment is consistent with what currently exists in Florida offshore waters to- day. According to Clench and Tumer (1950), both C dalli and C. pilsbr yi can currently be found in deep water off the Florida coast. Genus Opalia H. and A. Adams, 1853 Type Species: by subsequent designation, Scalaria australis Lamarck, 1822. Subgenus Nodiscala de Boury, 1889 Opalia politesae new species (Figures 29-33) Description: Shell small, slender; 2.5 smooth proto- conch whorls, 8 convex teleoconch whorls. Spire angle 20°. Twelve to 13 rounded axial costae on last teleoconch whorl. Shell surface covered with fine horizontal and vertical threads giving surface a pitted appearance. Pitted surface absent on distal surface of axial costae. Costae terminate at the suture, creating a crenulated ridge. Su- tures moderately impressed. No basal ridge present. No varices. Ape rture oblique, subcircular, ioe d by thick rounded lip. Inner portion of lip encircled by a thin, unsculptured rim. Outer portion possesses fine ‘vertical threads that radiate outwards towards periphery. No um- bilicus. Columella short and arched. Holotype: UF 114913, maximum width 1.7 mm. Type locality: Tenmile Creek 03 (CA017) (= Tulane University locality TU 546), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation. Paratypes: USNM 534495, USNM 534496 and UF 113897, 1 shell each, protoconchs missing, locality and formation for each specimen the same as holotype: USNM 534497, 1 shell, protoconch missing, Tenmile maximum height 5.5 mm, Page 116 THE NAUTILUS, Vol. 121, No. 3 Figures 29-38. Opalia politesac new species and Opalia mica new species. 29-32. Opalia politesac; apertural, lateral, abapertural, and basal views of holotype UF 114913; maximum height 5.5 mm, maximum width 1.7 mm, 33. Magnified view of sculpture of teleoconch of UF 114913. Scale bar = 0.41 mm, 34-37. Opalia mica; apertural, lateral, abapertural, and basal views ol holotype UF 66077; maximum height 4.0 mm, maximum width 1.5 mm, 38. Magnified view of sculpture of teleoconch of UF 66077. Seale bar () { boy G. W. Schmelz and R. W. Portell, 2007 Page 117 Creek 01 (CA002) (= Tulane University locality TU 830) Clarksville Quadrangle USGS 7.5! Series (1945), | ee County, Florida, C Jhipola Bonaaten: UF 67499, : shells, C dhipol: LOL (MecClelland’s Farm) (CA001) (= Tu lane Univ ersity locality TU 457), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation; UF 114914, 2 shells, locality and formation same as holotype; UF 114922, 3 shells, Ten- mile Creek 04 (CA020) (= Tulane University locality TU 951), Clarksville Quadrangle USGS 7.5’ Series (1945), aaee County, Florida, Chipola Ronnavon Distribution: Although Opalia politesae is one of the more common epitoniids found in the Chipola Forma- tion, its distribution appears limited to the reef paleoen- vironments found along Tenmile Creek and the Chipola River. Etymology: Named for Greta Polites, an avid student and ballecor of Chipola and Shoal River formation fos- sils. Discussion: According to Clench and Turner (1950: 231) members of the genus Opalia have “Spiral sculpture usually of exce edingly fine incised threads which may or may not be finely pitted.” In addition, the genus also possesses relatively low, heavy costae, no aanbilions and a thick outer lip divided into a thin inner unsculptured ring and a thicker finely pitted one. In the subgenus Nodiscala the sutures are crenulated and the basal cord is absent. Gardner (1947) reported the subgenus Nodiscala from Eocene deposits in Australia. However, there is no evi- dence of the genus Opalia from the Eocene of the United States. Machel and Dockery (1984) reported a number of different species of Opalia from the lower Oligocene Mint Spring Formation of Mississippi. Ac- cording to Gardner (1947: 578), the subgenus Nodiscala is * ‘peculiarly eee of the Miocene of central Europe” and she also noted that several species have been found in the Pliocene of Italy. Landau et al. (2006) reported a number of fossil Nodiscala from a wide range of European localities. These fossils have been found in deposits that date from the early Miocene to the late Pleistocene. DuShane (1979) lists Opalia borealis as pos- sibly coming from Miocene deposits of the northeastern Pacific region. All of the re maining Opalia studied by DuShane (1979) came from Pliocene and Pleistocene formations. In the Caribbean, Maury (1917) reported a single specimen of Epitonium textuvestitum from Pliocene deposits in Santo Domingo (Dominican Repub- lic). The description of this species given by Maury 1917 7) clearly indicates that it belongs to the genus Opa- lia. Both Campbell ( 1993) and Petuch (1994) have as- signed this species to the genus Opalia. Campbell's specimen came from Pliocene deposits in Hampton, Vir- ginia and Petuch’s specimen was reported from the Pliocene deposits of the former APAC shell pits in Sa- rasota. Florida. Opalia politesae is one of the more abundant species scacchi. (Hoernes, of Epitoniidae in the Chipola Formation where it is a unique representative of this genus. In overall contour and shape it bears some resemblance to the fossil O. textuvestitum but it lacks varices and has fewer costae on the body whorl. Among Recent species O. politesae is most similar to Opalia burryi, which ‘ranges from south Florida through the West Indies to Trinidad. However, O. politesae is a much smaller and more slender species that has fewer body whorls (S versus 9-11) and fewer costae on the body whorls (12-13 versus 14) than its Recent counterpart. The close similarity between O. po- litesae and O. burryi suggests that O. polite sae is prob- ably the ancestral form of O. burryi. Opalia mica new species (Figures 34-38) Description: Shell short, stocky; one smooth proto- conch whorl present, four teleoconch whorls. Nine to ten costae on last teleoconch whorl. Spire angle 29°. Costae sharply angulated, forming a node at the periphery of the body whorl. Shell surface crossed with fine horizontal and spiral threads that give the surface a pitted appear- ance. Pitted sculpture covers entire surface of axial cos- tae, Costae terminate at the suture creating a crenulated ridge. Sutures moderately impressed. No basal ridge present. No varices. No umbilicus. Aperture oblique, subcircular, surrounded: by a thick lip. Inner a of lip encircled by thin, smooth rim. Outer part of lip pos- sesses fine vertical threads that radiate outward towards the periphery. Vertical threads on lip overlapped by fine threads that give the lip a pitted appearance. Columella short and arched. Holotype: UF 66077, maximum height 4.0 mm, maxi- mum width 1.5 mm. Type Locality: Shoal River Grotto (WLO004) (= Tu- lane University locality TU 69A), New Harmony Quad- rangle USGS 7.5’ Series (1987), Walton County, Florida, Shoal River Formation. Paratypes: UF 114924, I specimen, and UF 88160, 3 specimens, locality and formation same as holotype. Distribution: —Opalia mica is a rare species but thus far has only been obtained from the type locality. Name alludes to its diminutive size. Etymology: Discussion: Opalia mica is structurally very different from O. politesae. It is broader and sme ler than the Chipola Formation species and possesses sharply angu- lated costae. Gardner (1947) reported fragments of this species from the Shell Bluff location along the Shoal River in Walton County, Florida. The most comparable fossil form to O. mica is the Miocene species Opalia ct. 1856) reported from Austria (Nord- sieck, 1972). In size and overall form the two species are very similar; however, O. mica has deeper sutures, lacks rounded varices, and does not possess the sutural spiral cords present in the European taxon. Among extant forms O. mica is similar to O pumilio Page 118 THE NAUTILUS, Vol. 121, No. 3 morchiana Dall, 1889. Both the Chipola Formation and Recent species are small and have costae that are sharply angulated at the periphery of the body whorl. However, O. mica as varices, has a more acute spire angle (29° versus 37°), and only has four teleoconch whorls com- pared to seven for O. pumilio morchiana. Genus Epitonium Réding, 1798 Type Species: by subsequent designation, Turbo sca- laris Linnaeus, 1758. Subgenus Asperiscala de Boury, 1909 Epitonium virginiae (Maury, 1910) (Figures 39-46) Description: Shell small, attenuate; 3 smooth, glossy, protoconch whorls, 6-7 convex teleoconch whorls sepa- rated by deep sutures. Spire angle 35°. 5-9 costae on last teleoconch whorl. Costae prominent, blade-like, slightly oblique with coronate shoulders. Fine spiral threads on body whorl crossed by faint axial growth line. No varices. Umbilicus absent. No basal cord. Aperture oval. Holotype: PRI 3467 (formerly in Cornell University collection), maximum height 3.75 mm, maximum width 1.5 mm. Other Material Examined: UF 95695, 1 shell, Chipola 13 (CA027) (= Tulane University locality TU 458), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Blond. Chipola Formation; UF 89251, 4 shells, Chipola 28 (CA06G) (= Tulane Univer- sity locality TU 545), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation: UF 103784, 1 shell, Tenmile Creek 06 (CA023) (= Tulane Univ ersity locality TU 456), Clarks- ville Quadrangle USGS 7.5" Series oe Calhoun County, Florida, Chipola For ition; UF 72341, 1 shell, Shoal River Grotto (WL004) (= Tulane Univ se locality TU 69A), New Harmony Olenge USGS 7.5’ Series (1987), Walton County, Florida, Shoal River Formation. Type Location: Florida. Bailey's Ferry, Calhoun County, Distribution: —Epitonium virginiae is one of the more common species of Epitoniidae found in the Chipola Formation. It is also found in the Shoal River Formation where it appears to be extremely rare. A total of 59 speci- mens of E. virginiae were examined from ten ( thipola Formation collection sites in Calhoun and Liberty coun- ties. The only Shoal River example of E. virginiae was obtained from UF loc ality WI ree = TU 69A) in Walton County. Etymology: Unknown. Discussion: Members of the genus Epitonium are small, thin, generally slender turriculate shells with a wide range of different sculptured characteristics. Some have body whorls that are attached while others do not. \ basal ridge may be present or absent, the costae may be thin and blade -like or thick and rounded, and spire al sculpturing may or may not be present. This high degree of structural variability within the genus has Caused: re- searchers to organize its members into numerous sub- genera. Epitoniidae with blade-like costae, spiral thread-like cords on the body whorls, and an absence of a basal ridge were assigned to the subgenus Asperiscala by de Boury (1909). Members of the subgenus Asperisc cala have been reported from deposits as old as Cretaceous (Wade, 1926). Clench and Turner (1952) kept the subgenus As- periscala and assigned to it all epitoniids with spiral cords and either blade or cord-like costae. DuShane (1979: 91), because of “Certain morphological differences from those of Epitonium s.s.”, elevated Asperiscala to full ge- neric rank when she described the family Epitoniidae’ in the northeastern Pacific. According to ‘Kilburn (1985), however, the type species of Asperiscala is not represen- tative for this taxon. Kilburn reported that the type spe- cies of Asperiscala described by de Boury (1909) had cancellate sculpture. abe, Kilburn assigned epitoniids with only spiral sculpture to the subgenus Parviscala. Weil et al. (1999) and Nakayama (2003) re- tained Asperiscala as a subgenus but limited its use to epitoniids that resembled Parviscala that have an open umbilicus and sutures with peaked costae. Herein, we have retained the use of Asperiscala in the sense of Clench and Turner (1952), pending resolution of the previo cited above. But unlike its use in Weil et al. and Nakayama, Asperiscala is herein used to represent mem- ees of the genus Epitonium that have nee sculpture that may or may not be intersected with faint axial threads or narrow cords. A more detailed analysis of larger specimens of E. virginiae revealed the presence of fine vertical lines that intersect the slender spiral threads on the body whorls (Figure 46). The absence of this feature in Maury’s de- scription was more than likely due to the small size of the specimen she examined. Of all the Chipola Formation species of Epitonium examined, only EF. virginiae was encountered in the younger Shoal River Formation. It is apparently a rare species in this unit given that numerous collecting trips to several different Shoal River Formation sites by dif- ferent collectors have yielded liane one specimen. Close examination of this specimen revealed that its shell struc- ture is fundamentally the same as that of the Chipola Formation species. Several fossil species similar to E. virginiae have been collected from different localities including the Carib- bean, Central and South America, and Spain. Epitoniwm (Asperiscala) venezuclense (Weisbord, 1962) from the upper Miocene Mare Formation of northern Venezuela comes closest in overall morphological characteristics to E. virginiae, Both species have prominent blade-like, slightly oblique costae with coroneted shoulders, and both have spiral threads in the interspaces between the axial costae that are crossed by fine axial filaments. The G. W. Schmelz and R. W. Portell, 2007 Figures 39-46. Epitonium virginiae (Maury, 1910). 39-40. Apertural and abapertural views of holotype PRI 3467; maximum ight 3.75 mm, maximum width 1.5 mm. 41. Magnified view of sculpture of teleoconch of PRI 3467. Scale bar = 1.0 mm, 42-45. \pertur il, lateral, abapt rtural, and basal views of UF 95695; maximum height 6.9 mm, maximum width 2.45 mm. 46. Magnified vi ot sculpture of teleoconch of UF 95695. Scale bar = 0.55 mm Page 120 THE NAUTILUS, Vol. 121, No. 3 two species differ from one another in that E. virginiae has more impressed sutures, the costae of E. virginiae do not become obsolescent near the posterior suture, and the angle of the spire is slightly wider in E. virginiae (35° versus 32°). Other fossil species similar to E. virginiae include Epi- tonium loripanum Pilsbry and Olsson, 1941, from the Pliocene of Ecuador, Epitonium amosbrowni. Pilsbry, 1921. from the Miocene of the Dominican Republic, Epitonium ct. gabbi (de Boury) Woodring, 1959, from the Gatun Miocene of Panama (Canal Zone), and Epi- tonium muiricatoides (Sacco, 1891) from the early Pliocene of Spain. The Chipola Formation species di ffers from E. amosbrowni, which has 16 axial costae on the last whorl versus 8-9 for E. virginiae, from E. cf. gabbi, which has more pronounced and numerous spiral threads on its body whorls, and E. muiricatoides, which lacks spiral cords on the body whorls. A close comparison of E. loripanum with E. virginiae suggests that they may be the same species. Bok have the s same body shape, the presence of faint spiral threads on the body whorls, the same number of teleoconch whorls, thin ribs with coronated shoulders, and lack an umbilicus. Among extant species, E. virginiae is most similar to Epitonium denticulatum (Sowerby, 1844). However, E. virginiae is more slender and has fewer axial costae. Epitonium incomitatum new species (Figures 47-51) Description: Shell small, thin, turriculate. Protoconch missing; 6 strongly convex teleoconch whorls separated by deep sutures. Spire angle 31°. Thin, low, reflected costae on last teleoconch whorl number ing 24. Costae on each succeeding whorl slightly offset from the one above. Numerous spiral cords on body whorl. No varices. No basal ridge. Umbilicus present. Outer lip of aperture thin. Aperture suboval. Holotype: UF 91452, maximum height 5.0 mm, maxi- mum width 2.2 mim. Tenmile Creek 03 (CAQ17) (= Tulane Clarksville Ouadenel Calhoun County, Florida, Type Locality: University locality TU 546), USGS 7.5' Series (1945), ¢ Chipola Formation. Distribution: Only known from the type locality. Etymology: The name is derived from the Latin word incomitatus meaning unaccompanied or alone. It refers to the unique specimen (holotype). Discussion: = Epitonium incomitatum is an extreme ly fragile shell. In shell sculpture it is similar to Parvise ala, a subgenus retained by Weil et al. (1999) and N: tkayame U 2003). Gardner (1947) re porte da fragment of this shell in the ( thipola Formation and assigned it to the subgenus Crisposcala. A fragment similar to E. incomitatum was also reported from the upper part of Gatun Formation in Panama (Canal Zone). Woodring (1959) tentatively iden- tified this species as Epitonium rushii (Dall, 1889). Weil et al. listed E. rushii as synonym of Epitonium striatis- simum Monterosato, 1878. Among fossil western Euro- pean species E. incomitatum is most similar to Epito- nium pulchellum (Bivona, 1832) which has been re- ported from the middle Miocene in Italy (Cavallo and Repetto, 1992). Epitonium incomitatum differs from its European counterpart by having less elevated varices and less prominent spiral cords on the body whorls. Among extant species, E. incomitatum is most similar to E. striatissimum, a rare species found in shallow water off Cape Hatteras. se ae incomitatum differs from E. striatissimum in that the body whorls are more in- flated (the angle formed with the spire is 31° versus 25 for E. striatissimum) and the spiral cords on the body whorl are much broader and less numerous. Epitonium incomitatum also bears some resemblance to Epitonium multistriatum Say, 1526, a species that ranges from Massachusetts to Texas. It differs from this species by ie: more numerous costae on the last body whorl (25 versus 16-19) and by having broader spiral cords on the body whorls. In addition, in E. in- comitatum the costae are less abundant in the early whorls, while E. multistriatum the costae are more nu- merous in the early whorls (as many as 43 in some speci- mens). Epitonium regina new species (Figures 52-56) Description: Shell small, turriculate: three smooth, glossy protoconch whorls, eight slightly angular body whorls separated by a moderately deep suture. Spire angle 23°. Eleven narrow, low, slightly reflected costae on last teleoconch whorl. Costae occasionally offset with costae on preceding whorl. Numerous faint, spiral cords on body whorls intersected by faint spiral threads. No basal cord. No umbilicus. No varices. Aperture thin, sub- OV. al. Holotype: USNM 534487, maximum height 6.1 mm, maximum width 2.1 mm. Farley Creek 03 (CAO009) (= Tulane Clarksville Dlsdmel Calhoun County, Florida, Type Locality: University locality TU 825), USGS 7.5’ Series (1945), Chipola Formation, Paratype: UF 67498, I specimen, Chipola OL (CAOOL) (McClelland’s Farm) (= Tulane University locality TU 457), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun C ounty, F lorida, € Chipola Formation. Distribution: Collected only from the type locality. Etymology: Named from the Latin word regina, meaning queen, an allusion to the stately appearance of this species. Discussion: regina bears a slight resemblance to Epitonium obliquum (Sowerby, 1$47), however E. regina has a cancellate pattern be- Among Recent species, E. G. W. Schmelz and R. W. Portell, 2007 Page 121 Figures 47-56. Epitoniwim incomitatiwm new species and Epitonium regina new species. 47-50. Epitonium incomitatum, ape tural, lateral, abapertural, and basal views of holotype UF 91452; maximum height 5.0 mm, maximum width 2.2 mm. 51. Magnified view of sculpture of teleoconch of UF 91452. Scale bar = 0.41 mm. 52-55. Epitonium regina: apertural, lateral, abapertural, and basal vs of holotype USNM 534487: maximum height 6.1 mm, maximum width 2.1 mm. 56. Magnified view of s¢ ulpture of teleoconch of USNM 5344587. Scale bar = 0.65 mm Page 122 THE NAUTILUS, Vol. 121, No. 3 tween costae on the body whorls, lacks elevated costae near the sutures, and has no umbilicus. Among fossil species E. regina bares some resem- blance to Epitoniwm smithfie Ider sis Mansfield, 1929 and Epitonium dupliniana (Olsson, 1916). Epitonium smith- fieldensis was reported from the Pliocene Yorktown For- mation of Virginia. Like the Chipola Formation species it is ornamented with marginally reflected slender varices and the number of varices on the last body whorl is 12. However, E. smithfieldensis does not have any spiral sculpture and its varices are united at the suture. Epito- nium dupliniana was reported from the middle Pliocene Duplin Formation of North Carolina (Olsson, 1916). Like E. regina, it has a thin shell with low varices and spiral sculpturing on the body whorls that is intersected by vertical threads. It differs from the Chipola Formation species in that the varices on E. regina are broader and more cord-like than they are in E. dupliniana. Also, the upper shoulder on each of the varices of E. dupliniana has a small hook-like projection, a feature not et on the varices of E. regina. Subgenus Epitonium Réding, 1798 Type of Subgenus: = Tubo scalaris Linnaeus, 1758 by subsequent designation, Suter, 1913. Epitonium contwaide new species (Figures 57-66) Description: Shell small, attenuate; 3.5 smooth proto- conch whorls. Spire angle 30°. Six-and-a-half moderately convex teleoconch whorls separated by a depressed su- ture. Seven to nine low, moderately broad, T-shaped costae on last body whorl. Costae gradually increase in width as aperture ‘approache .d. Costae angled at shoul- ders. Body whorls smooth but eroded specimens exhibit numerous, fairly broad spiral cords on body whorls. No basal cord, umbilicus, or varices Aperture suboval, outer lip slightly expanded and thicketie d. Holotype: UF 113894, maximum height 3.2. mm, maximum width 1.2 mm. Type Locality: Shoal River Grotto (WLO04) (= Tu- lane University locality TU 69A), New Harmony Quad- rangle USGS 7.5’ Series (1987), Walton County, Florida, Shoal River Formation. Paratypes: UF 67208, 9 shells, Shell Bluff ( (WL002) (= Tulane University locality TU 69), New ae mony Quadrangle USGS 7.5’ Series (1987), Walton County, Florida, Shoal River Formation; UF 89549, 1 same as preceding; UF 117092, 2 shells, locality and formation same as preced- shell, locality and formation s ing: UF $9638, 29 she lls, locality and formation same as preceding; UF 72340, 1 shell, locality and formation same as holotype; UF 88170 locality and formation same as holotype Distribution: Epitonium contwaiae is known only from the type locality and from the Shell Bluff on Shoal River. Etymology: Named for Wendy Conway a long time field associate of the authors and an avid eallector of Chipola and Shoal River formation fossils. Discussion: —Epitoniwm with and without an umbilicus and possessing smooth body whorls and no basal cord were assigned to the subgenus Epitonium by Clench and Turner (1951). Later researchers. including Weil et al. (1999) and Nakayama (2003) subdivided Epitonium with smooth body whorls into a number of subgenera. For example, t those with smooth body whorls, no umbilicus, and peaked costae were assigned to the subgenus Hir- toscala, while Epitonium \y with smooth body whorls, peaked costae and a slit-like umbilicus were placed into the subgenus Lamelliscala. Additionally, those with smooth body whorls, no umbilicus, and thick costae were assigned to the subgenus Nitidiscala. Although probably in the minority, we find the need for splitting out the genus Epitonium into so many ad- ditional subgenera based upon minor morphological dif- ferences probably counterproductive to the establish- ment of a more realistic classification system. Clench and Tumer (1951) wamed researchers about this problem when they reported the difficulties de Boury experienced with his attempts to split out different members of the family. Rather than add to the confusion that exists with this taxonomic group, we have reverted to using the broader subgeneric name of Epitonium as defined by Clench and Turner (1951). When compared to Recent species, non-eroded speci- mens of Epitonium conwaiae (Figures 57-61) are most similar to Epitonium humphreysii Kiener, 1838, which ranges from Cape Cod Massachusetts south to Florida (excluding the Florida Keys) and into the Gulf of Mexico from Cape Romano to Texas. Fossil specimens of E. humphreysii have also been reported from upper Mi- ocene deposits of the Entrerriense Formation of the Chubut Province, Argentina (Brunet, 1995). Both the Shoal River Formation species and its Recent and fossil counterpart have flattened costae which are variable in width, both lack sculpturing on the body whorls, and each species has a similar number of costae on the last body whorl. The Shoal River Formation species differs from E. humphreysii by having more deeply impressed sutures, more angular costae on the shoulder of the body whorls, and much smaller ave rage size. In eroded specimens of E. conwaiae (Figures 62-66), the body whorls reveal a sculpture of broad spiral cords most similar to Epitonium championi Clench and Turner, 1952, a rare epitoniid that inhabits intertidal and near-shore waters from Cape Cod to North Carolina. Both the Shoal River Formation form and E. championi have flattened, cord-like costae which are variable in width and both have spiral sculpturing which consists of numerous flattened spiral cords. The Shoal River For- mation species, however, differs from E. championi by having more deeply impressed sutures, a greater number of costae on the last body whorl (11 versus S or 9) and a G. W. Schmelz and R. W. Portell, 2007 Figures 57-66. = Epitonium conwaiae new species. 57-60. Apertural, lateral, abapertural, and basal views of holotype UF 113594; iaximum height 3.2 mm, maximum width 1.2 mm. 61. Magnified view of sculpture of teleoconch of UF 113894. Scale bar = 0.7 mm. 62-65. Apertural, lateral, abapertural, and basal views of paratype UF 72340; maximum height 4.2 mm, maximum width 2.0 61. Magnified view of s« ulpture of teleoconch of UF 723 AQ. Scale bar = 1.26 mm. UF 72340 is an eroded specimen figured here to compare sculptural differences between it and unworn holotype UF 113894 greater angle between the spire and the outer shoulders of the shell (30° versus 20 Among fossil species, well-preserved specimens of E conwwaiae are most similar to Epitonium boltoni Gardner 1948, from the Pliocene Tar River deposits in North Carolina Like E. conwaiae, E boltoni possesses smooth spiral whorls, has thickened slightly raised costae an lacks an umbilicus and basal cord. However, E. conwaia differs from E. boltoni by having more impressed su tures, more angular costae on the dorsal surface of each body whorl and fewer numbers of costae on the last bods whorl (9 versus 12 Page 124 THE NAUTILUS, Vol. 121, No. 3 The eroded form of FE. conwaiae also bears some simi- larities to Epitonium alaquaense collected and described by Mansfield (1935) from the late Miocene strata of Vaughan Creek in Walton County, Florida. Gardner (1947: 577) reported collecting a partial specimen from Walton County from “a horizon that was slightly higher than the typical Shoal River formation.” These i investiga- tors were not able to locate Gardner's shell fragment but were able to borrow Mansfield’s holotype (USNM 373149) for comparative purposes (see Figures 67-68). Unfortunately, the holotype was broken, which made the comparative study a bit more difficult than expected. Nevertheless, examination of the shells showed that E. conwaiae differs in several ways from E. alaquaense. Epi- tonium conwaiae is more slender than E. alaquaense and has 3.5 nuclear whorls versus 2.5 for E. alaquaense and the number axial ribs on the post-nuclear whorls on E. conwaiae range from 7-9 while the number of axial ribs on E. alaquaense ranges from 9-12. Current evidence suggests that that E. alaquaense is not found in the younger Shoal River Formation since none of the speci- mens examined were similar to Manstield’s shell. The eroded form of E. conwaiae is also similar to the fossil species Epitonium santodomingonum Pilsbry, 1921, from the Pliocene beds of Santo Domingo (Do- minican Republic) and Epitonium antillarum fe Boury, 1909) from Pliocene beds in Virginia and North Carolina (Gardner, 1948) and Florida (Olsson and Harbison, 1953). Like E. conwaiae, both fossil Pliocene species pos- sess numerous spiral cords on the body whorls, lack a basal ridge, and both have low, well de veloped costae on the body whorls which increase in width near the aper- ture. Epitonium conwaiae differs from E. santodomin- gonum by having fewer costae on the last body whorl (9 versus 18) and an absence of varices on the last whorl of the shell. Epitonium antillarum differs from E. conwaiae by possessing fewer varices (7-9 versus 10-13), fewer teleoconch whorls (6 versus S—9), thinner costae on the body whorls, a thinner lip surrounding the aperture, and a more acute spire angle (22° versus 30°). It should be noted that E. antillarum is no longer considered a valid species. In 1909, de Boury assigned the species antil- larum to the epitoniid Sc alaria turricula Sowe rby, 1844. However, the shell Sowerby (1844) described had al- ready been named by 7’ nee (1842) as Sealaria can- deana. Subsequently, Clench and Turner (1952) rectified de Boury’s mistake by recognizing E. antillarum as Epi- tonium candeanum. Of Favithe yr note, the shell described by Clench and Turner (1952) as E. candeanum does not fit the description of E. antillarum given by Gardner 1948) and later listed by Olsson and Harbison (1953) in their treatise on Pliocene Mollusca of Southern Florida. \ccording to Clench and Turner (1952), E. candeanum has thinner and more numerous costae (18-25 versus 10-13) on the body whorls than does the species de- scribed by Gardner (1945) as £. antillarum. In all prob- ability the Epitonium described by Gardner (1948) is a new spe cies. It is beyond the scope of this paper to rec- tify this error and any effort to do so is being left to the work of future investigators. Epitonium hoerleae new species (Figures 69-73) Description: Shell medium height, sturdy, turriculate; 3.5 smooth, glossy, protoconch whorls. 7.5 convex teleo- conch wars. separated by deep suture. Spire angle 24°. Eight to nine thin costae on last body whorl. ‘Costae slightly reflected backwards. At the whorl shoulder cos- tae are slightly expanded and form a cusp-like node. Cos- tae connected to one another at the suture, forming an oblique angle to the shell’s central axis. Extremely ant spiral thivends on glossy body whorls. No umbilicus. No basal cord. Aperture suboval. Outer lip of aperture thin and reflected backwards. Holotype: USNM 534488, maximum height 8.8 mm, maximum width 3.2 mm. Type locality: Tenmile Creek 03 (CAQ17) ( University locality TU 546), USGS 7.5’ Series (1945), Chipola Formation. Paratypes: USNM 534498, 1 shell, locality and forma- tion same as holotype: UF 84579, protoconch missing and aperture broken, Tenmile Creek 01 (CA002) (= Tu- lane University locality TU 830), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation; UF 101862, 1 shell, protoconch missing and aperture broken, Tenmile Creek 13 (CAQ58) (= Tulane University locality TU 1097), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipota Formation; UF 99083, 1 shell, Chipola 23 (CA037) (= Tulane University locality TU 711), Clarksville ae USGS 7.5’ Series (1945), Cal- houn County, Blonde: C eae Formation; UF 94317, 2: shells, Chipola 13 (CA027) (= Tulane University locality TU 458), Clarksville Gundiancle € USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation. = Tulane Clarksville Quadrangle Calhoan County, Florida, Distribution: Epitonium hoerleae is a ve ry common, widely distributed species. It has been found at the type locality of Tenmile Creek as well as along F Farley Creek and the Chipola River. Etymology: The species was named in honor of Shir- ley Hoerle, one of the pioneer re searchers of C hipola Formation mollusks. Discussion: Clench and Turner (1951) placed e Saat niids that possessed glossy whorls and extremely faint spiral threads into the subge snus Epitonium. This classi- fication has been retained with E. hoerleace. Among recent species E. hoerleae is most similar to Epitonium foliaceicostum VOrbigny, 1842, which inhab- its offshore waters in the Caribbean and along Florida's east coast. Both the fossil species and its Recent coun- terpart have glossy body whorls with faint spiral threads, approximately the same number of costae on the last body whorl (S—9 versus 7—S), and a slightly reflected G. W. Schmelz and R. W. Portell, 2007 Page 125 Figures 67-73. Epitonium alaquaense Mansfield, 1935, and Epitonium hoerleae new species. 67-68. Epitonium alaquaens« ipertural and abapertural views of holotype USNM 373149 shown for comparison to E. conwaiae. 69-72. Epitonium hoerl ypertural, lateral, abapertural, and basal views of holotype USNM 534488; maximum height $.8 mm maximum width 3.2 mm Maenified view of s« ulpture of teleoconch of USNM 534488. Scale bar 0.95 mm a i 2 od. Page 126 THE NAUTILUS, Vol. 121, No. 3 aperture lip. Epitonium hoerleae differs from the Recent shell by having a greater number of nuclear whorls (3.5 versus 1.5), less alee ated costae and a more slender body configuration. A comparison of E. hoerleae with a number of fossil species shows that it most closely resembles Epitonium fargoi Olsson and Harbison, 1953, which was described from the Plio-Pleistocene Caloosahatchee Formation of south Florida as well as Epitonium proximus (de Boury, 1890) which was described from the early Pliocene of France and the middle Pliocene of England (Harmer, 1920-1925). Both E. hoerleae and E. fargoi have convex, glossy, body whorls, approximately the same number of costae on the last body whorl (§ versus 9), the costae are reflected backwards, and both have a small cusp-like node on the upper shoulder. Epitonium hoerleae differs from E. fargoi in that it has a more expanded reflected outer lip, lacks a thickened cord on the inner lip, its costae are thinner and more blade-like and the spire angle in E. hoerleae is less acute (24° versus 20°) than its Pliocene counterpart. The European species, E. proxi- mus, differs from E. hoerleae in that it is larger and more slender than its American counterpart and has more va- rices on the body whorl (11-13 versus S—9). Epitonium kallistos new species (Figures 74-78) Description: Shell small, turriculate; 3.5 smooth, glossy, protoconch whorls, 6.5 convex teleoconch whorls separated by a deep suture. Six moderately thickened, slightly recurved costae on last body whorl. Spire angle 25°. Costae connected to one another at suture forming an obliq ue angle with the central axis. Costae lack a cusp- like bode on the shoulder of body whorl. Body whorls smooth and glossy. No umbilicus; basal cord absent. No varices. Aperture suboval, lip slightly thickened and re- curved. Holotype: UF 44614, maximum height 6.8 mm, maxi- mum width 2.3 mm. Type Locality: Tenmile Creek 02 (CA003) (= Tulane University locality TU 70), Altha West Quadrangle USGS 7.5’ Series (1982), Calhoun County, Florida, Chipola Formation. Distribution: the type locality. Epitonium kallistos is known only from Etymology: Name is derived from the Greek word kallisto meaning most beautiful. Discussion: Among Recent and fossil epitoniids, E kallistos is most similar to Epitonium unifasciatum (Sow- erby, 1844), a species that today r anges from southern Florida to the Lesser Antilles (Clench and Turner, 1951) Epitonium unifasciatum has also been reported from the upper Miocene deposits of the Entrerriense Formation Argentina (Brunet, 1995). Like its Recent and fossil counte rpart, E kallistos has smooth, of Chubut Province shiny, convex whorls, the aperture 1s suboval, there is no basal ridge, and the outer lip of the aperture is reflected backwards. Epitonium kallistos differs from E. unifascia- tum by having fewer costae on the last body whorl (6 versus 7 — 9), a greater spire angle (25° versus 20°), more deeply impressed sutures, and the costae on the body whorls are more elevated. Subgenus Gyroscala de Boury, 1887 Epitonium vokesae new species (Figures 79-83) Description: Shell small, turriculate; three smooth, glossy, bulbous protoconch whorls, 5.5 moderately con- vex ieee whorls. Suture moderately impressed. Spire angle 27°. Eight to nine slightly raised, blade-like costae on boty whorl. Costae sinuous, not joined at su- ture with costae on preceding whorl. Distinct, narrow, basal ridge present. No varices. No umbilicus. Outer lip missing. Holotype: UF 113595, maximum height 4.3 mm, maximum width 1.6 mim. Type Locality: Tenmile Creek 04 (CA020) (= Tulane University locality TU 951), Clarksville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chipola Formation. Paratypes: UF 114957, 1 shell, Tenmile Creek 03 (CAO017) (= Tulane University locality TU 546), Clarks- ville Quadrangle USGS 7.5’ Series (1945), Calhoun County, Florida, Chapels haiti, UF 99136, 1 shell, spire missing, Chipola 23 (CAQ37) (= Tulane University locality TU 711), Clarksville Quadrangle USGS 7.5’ Se- ries (1945), Calhoun County, Florida, Chipola Forma- tion. Distribution: Epitonium vokesae has only been col- lected along Tenmile Creek and the Chipola River. Etymology: Named in honor of Dr. Emily Vokes, a leading researcher of the family Muricidae and a mentor to the many who have studied Chipola Formation fossils. Discussion: Epitonium of the subgenus Gyroscala possess a smooth protoconch of about three whorls, high axial lamellae, gently convex smooth body whorls, a well defined, thin basal cord and a thickened peristome (Kil- burn, 1985). Thiele (1929) and a number of other European work- ers have made Gyroscala subordinate to Cirsotrema. Clench and Turner (1951) and Abbott (1974) retained de Boury’s (1887) taxonomic classification of Gyroscala as a subgenus of Epitonium. Kilburn (1985) followed the lead of Australian and Japanese malacologists and accorded Gyroscala full generic status. According to Nakayama (2003), the presence of a basal cord in Gyrose ala justifie s raising this epitoniid to generic level. Once Gyroscala was raised to generic status, a number of investigators suggested establishing subgenera. Kilburn, for example, suggested two subgenera (Boreoscala and Circuloscala), while Nakayama suggested three (Fragiliscala, Pomi- scala, and Circuloscala). As noted earlier, Nakayama al- G. W. Schmelz and R. W. Portell, 2007 Page 127 Figures 74-83. Epitonium kallistos new species and Epitonium vokesae new species 74-77. Epitonium kallistos apertural lateral ibapertural, and basal views of holotype UF 44614; maximum height 6.8 mm, maximum width 2.3 mm. 78. Magnified view of sculpture of teleoconch of UF 44614. Scale bar = 0.51 mm 79-82. Epitonium vokesat apertural lateral abapertural and basal of holotype UF 113898; maximum height 4.3 mm, maximum width 1.6 mm. 83. Magnified view olf sculpture of teleoconch of UI 113898. Scale bar = 0.37 mm Page 125 THE NAUTILUS, Vol. 121, No. 3 ready used Boreoscala as a subgenus for Cirsotrema. To further complicate the issue Weil et al. (1999) elevated Gyroscala to the generic level but avoided using any subgenera. Until a more comprehensive study has ‘Ge en done with Gyroscala, we have decided to retain Gyro- scala as a subgenus of Epitonium as suggested by de Boury and retained by Clench and Turner. Representativ es of the subgeneric group Gyroscala are not very common. There are three reported living species of the subgenus Gyroscala in the western Atlan- tic. Epitonium late llosum Lamarck, 1822, is found from Lake Worth, Florida, to the Lesser Antilles, as well as from France to South Africa. Recently, Garcia (2002), reported E. lamellosum from the Indo-Pacific to Califor- nia. Epitonium rupicola Kurtz, 1860, is found from Prov- incetown, Massachusetts, in the Atlantic to the Texas coast in the Gulf of Mexico, whereas Epitonium xenicima Melville and Standen, 1903, is a circum-global species (Garcia, 2006). None of the Recent western Atlantic spe- cies appear similar to the Chipola Formation species which is much more slender, lacks varices on the last whorl, has far fewer costae on the body whorls, and pos- sesses a very distinctive, bulbous protoconch. There are three fossil species similar to E. vokesae. One is Epitonium aciculum (H. C. Lea, 1843) which, according to Campbell (1993), was misidentified by Gardner (1948) as E. pratti. The species comes from the Pliocene Yorktown Formation in Virginia and North Carolina as well as the Plio-Pleistocene Waccamaw For- mation in North Carolina. Epitonium vokesae is more slender than the Virginia and North Carolina species. In addition, it has far fewer costae on the body whorl (9 versus 16-25), The other two fossil species similar to E. vokesae are an unnamed specimen from the Miocene Chagres Formation of Panama (Woodring, 1959) and Epitonium magnolianum (Olsson, 1916) from the lower to middle Pliocene deposits from North and South Caro- lina and Georgia. Both the Chipola Formation and Panama fossil species lack spiral sculpturing, have low sinuous costae on the body whorls, and a suppressed basal disk. The Chipola Formation species differs from the Panama fossil in that it is much more slender and has far fewer costae on the body whorl (9 versus 21). Epito- nium vokesae is also a much more slender species than E. magnolianum and lacks varices. ACKNOWLEDGMENTS We extend a special note of thanks to Burt Hayes, Cecil Sexton, and William Tatum (Calhoun County, Florida) and Robert Larson (Walton County, Florida) for allowing access to collect on their properties, Thomas Waller and Warren Blow (USNM), Bushra Hussani (AMNH), Paul Callomon (ANSP), Warren Allmon and Jonathan Hen- dricks (PRI), and Gustav Paulay and John Slapcinsky FLMNH provided specimen loans or access to collee- tions under their care Jose H. | eal (BMSM) kindly pro- vided digital images of paratype BMSM 15301 (C. cirri- tum). Andy Murray (Bradenton, FL), Greta Polites (Uni- versity of Georgia, ee Kevin Schindler (Flagstaff, AZ), and W ends Conway (Sun City, FL) helper with specimen donations and field assistance. Sean Roberts (FLMNH) assisted with digital photography using a Sony DSC RI camera (10.3 megapixel resolution). ). Support for field studies was provided by the McGinty Endowment of the Florida Museum of Natural History and by Bar- bara and Reed Toomey (both to R.W.P.). Thanks to the anonymous reviewers who improved the final version of the manuscript. This is University of Florida Contribu- tion to Paleobiology 595. LITERATURE CITED Abbott, R. T. 1974. American Seashells. 2" edition. Van Nos- trand-Reinhold, New York, 663 pp., 24 pls. Brunet, R. F. J. 1995. New species of Mollusca from the En- trerriense Formation (Upper Miocene) of Chubut Prov- ince, Argentina and species not previously reported from this Formation. Part 1—Gastropoda and Scaphopoda. Tu- lane aes *s in Geology and Paleontology 28: 1-56, 5 pls. Campbell, L. D. 1993. Phocene Molluscs from the Yorktown a ares River formations in ae Virginia Divi- sion of vee ral Resources Publication 127: 171 pp., 43 pls. Carter, |. G., T. J. Rossbach, Z. P. Mateo, and M. J. 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Geology and Paleontology of the Canal Zone and adjoining parts of Panama. United States Geo- logical Survey Professional Paper 306-B: iii + 147-239, pls. 24-38, THE NAUTILUS 121(3):131-138, 2007 Page 13] Molecular phylogeny of some Indo-Pacific genera in the subfamily eee H. Adams and A. Adame. 1853 (1838) (Gastropoda: Neogastropoda) Francisco M. Heralde HI National Institute of Molecular Biology and Biotechnology Maren Watkins Marine Science Institute University of the Philippines Diliman, Quezon City PHILIPPINES Gisela P. Concepcion Marine Science Institute University of the Philippines Diliman, Quezon City PHILIPPINES University of Utah John-Paul Ownby Pradip K. Bandyopadhyay and Department of Biology University of Utah Salt Lake City, UT 84112 USA Ameurfina D. Santos National Institute of Molecular Biology and Biotechnology : University of the Philippines Diliman, Quezon City PHILIPPINES Baldomero M. Olivera! Department of Biology Salt Lake City, UT 841 12 USA olivera@biology.utah.edu ABSTRACT We have carried out a phylogenetic analysis of a group of Indo- Pacific species in the subfamily Turrinae (Swainson, 1840) us- ing 12S mitochondrial ribosomal RNA gene sequences. Most of the species analyzed are convention: ally assigned to a: of three genera, Turris Roding, 1798, Gemmula Weinkauff, 1875 or Lophiotoma Casey, 1904. The molecular analysis ers that while the species of Turris and Gemmuila analyzed in this study comprise monophyletic groups, the species presently assigned to Lophiotoma definitely do not constitute a monophyle tic as- semblage and can be separated into two very distinctive groups of species based on the molecular analysis. The species pres- ently designated as Lophiotoma tayabasensis Olivera, 2004, Lophiotoma panglaoensis Olivera, 2004, Lophiotoma indica (Réding, 1798) and Lophiotoma bisaya Olive pra, 2004, are re- lated to Lophiotoma | (Une ‘dogemmula) unedo (Kiener, 1839 in 1S34-S0) by molecular criteria, and are clearly ence ly sepa- rated from Turris, Gemmula or L ophiotoma (as redefined). We propose that Unedogemmula (MacNeil, 1960) be recognized as a full genus; Unedogemmula unedo (Kiener, 1839 in 1834—S0) is the type species, and the species above are transferred from Lophiotoma to Unedogemmula INTRODUCTION Venomous gastropods comprise three groups: the cone snails, the auger snails or terebrids and the Ba classically included in a single family, Turridae (H. Ad- ams and A. Adams, 1853): these families are ann assigned to the superfamily Conoidea (Ponder and ' Author for correspondence Waren, 1988: Taylor, Kantor, and Sysoev, 1993). It has been apparent for some time that the * otainaele? comprise the largest species group in the superfamily (see for ex- ample, ‘Powell: 1966): however, the groundbre: iking work ot Bouchet and co-workers ( Bouchet et al., 2002: 2004) in New Caledonia, has provided evidence that >90% of Conoidean biodiversity probably resides in the “turrids” (broadly defined). “Turrids” are problematic at all levels: not only are they a megadiverse group (>10,000 species) with a large fraction of species that remain undescribed, but the phy- logenetic relationships within the group are poorly un- derstood. The number of different “turrid” genera that have been proposed is >600; although in traditional mol- luscan taxonomic work all turrids had been assigned to the family Turridae, in most of the more recent system- atic treatments, the group has been split into 5-6 ‘differ- ent families (Taylor et al., 1993; Bouchet and Rocroi, 2005). However, some standard taxonomic treatments retain the old nomenclature (see for example Kohn, 1998S). To complement our ongoing study of turrid venoms (see for example Watkins et al., 2006) we initiated « a study of “turrid” molecular phylogeny: the first results at the generic level are reported below. The genus Turris Réding, 1798, is the nominate genus of the family Tur- ridae, with Turris babylonia (Linnaeus, 1758) as the type species. This is an e xclusive ly Indo-pacific ge nus. How- ever, there has been inconsistency in the definition of those genera which are traditionally oe together with Turris in the subfamily Turrinae. Two other groups largely from the Indo-Pacific, Lophiotoma Weinkaut! 1875, and Gemimula Casey, 1904, are included in the Page 132 THE NAUTILUS, Vol. 121, No. : subfamily by most workers; in the western Atlantic aa eastern Pacific, the genus Polystira Woodring, 1928, also thought to have a close affinity to Turris (the es species for these genera are Lophiotoma acuta (Perry, 1811), Gemmula hindsiona (Berry, 1958) and Polystira albida (Perry, 1811)) Additionally, a number of other groups (such as Tur ridh upa (Hedley, 1922)) are regarded as turrine by most workers. A major motivation for these studies is to investigate the toxin genes expressed in the venom ducts of gastropods. Among the different groups of conoidean <¢ turrids, we have initially coucentrated on studying the gene products expressed i in the venom ducts of species in the subfamily Turrinae, since they are larger and more easily collected than are most other forad groups. We hope to correlate the gene families expressed in venom ducts with the nielecular phylogeny of the species ana- lyzed. Thus, the molecular analy sis reported below has focused on lar ger Indo-Pacific species in the subfamily Turrinae, e.g. Turris, Lophiotoma and Gemmula spp. The data that we present below demonstrates that two groups of species presently assigoned to the genus Lophiotoma, which appear to be dloscly related by shell morphology, prove to be unexpecte dly divergent when evaluated by molecular criteria and need to be placed in different genera. MATERIALS AND METHODS Specimen Collection: — Species analyzed in this study, shown in Table 1, were mostly collected by commercial shell collectors in the Philippines, except for Polystira albida (Perry, 1811), a generous gift of Drs, Estuardo Lopez Vera and Ed Heimer, and Lophiotoma cerithifor- mis (Powell, 1964), which was collected in Oahu, Hawaii. Specimens of each were preserved either in RNAlater® (Ambion Inc., Tx) or 95% ethanol, and DNA extracted as described below. In most cases, the digestive gland was used as the source of DNA; however, for alcohol pre- served specimens where the shell had not been cracked, the digestive gland was often degraded, and DNA was extracted from foot tissue. Identification and sequencing of clones encoding 12S mitochondrial rRNA gene segments: Genomic DNA was prepared from tissue (~20 mg) from each turrid species using the Gentra PUREGENE DNA isolation kit (Gen- tra Systems, Minneapolis, MN) according to the manu- renomic DNA from each species (~lOng) was use dias a template for polymerase fac ture rs standard protocol. ¢ chain reaction (PCR) with oligonucleotides correspond- ing to 12S-I (5' TGC CAG CAG YCG CGG TTA ) and 12S-III (5' AGA GYG RCG GGC GAT GTG T) mito- chondrial rRNA segments (Oliverio and Mariottini, 2001). The 5’ and 3’ primers included adapte rs GGAGA- CAU and GGGAAAGU respectively for annealing to the cloning vector pNEB206A. The POR cycling profiles vere as follows: initial denaturation (95°C, 60s): followed hy 40 cycles of denaturation (95°C, 20s): annealing ( 55°C. 20s) and extension (72°C, 30s), The PCR products were ee Table 1. List of species analyzed in this study. Species Locality Lophiotoma acuta (Perry, 1811) Lophiotoma bisaya® Olivera, 2004 Lophiotoma cerithiformis (Powell, 1964) Lophiotoma cingulifera (Lamarck, 1822) Lophiotoma indica® (Réding, 1798) Lophiotoma Jickelii (Weinkautf, 1875) Lophiotoma olangoensis Olivera, 2002 Lophiotoma panglacensis® Olivera, 2004 Lophiotoma polytropa (Helbling, 1779) Lophiotoma tayabasensis® Olivera, 2004 Lophiotoma unedo® (Kiener, 1839 in 1834-80) Gemmula speciosa (Reeve, 1543) Gemmula diomedea Powell, 1964 Gemmula rosario Shikama and Hayashi, 1977 Gemmula lisajoni Olivera, 2000 Gemmula sogodensis Olivera, 2005 Turris garnonsii (Reeve, 1$43) Turris grandis (Gray, 1833) Turris normandavidsoni Olivera, 2000 Turris babylonia (Linnaeus, 1758) Turris spectabilis (Reeve, 1S43) Turris totiphyllis Olivera, 2000 Polystira albida (Perry, 1811) Drillia regius (Habe and Murakami, 1970) Buenavista, Marinduque, Philippines Batangas, Philippines Oahu, Hawaii Cawoy, Olango Island, Philippine S Aligway Is. Dipolog Philippines Cawoy, Olango Island, Philippine S Cawoy, Olango Island, Philippines Panglao Is. Bohol, Philippines Bataan, Luzon, Philippines Sogod, Cebu, Philippines Panglao Is. Bohol, Philippines Batangas, Philippines Sogod, Cebu, Philippines Sogod, Cebu, Philippines Sogod, Cebu, Philippines Sogod, Cebu, Philippines Cawoy, Olango Is and, Philippines Sogod, Cebu, Philippines Sogod, Cebu, Philippines Cawoy, Olango Island, Philippines Cawoy, Olango Island, Philippines Cawoy, Olango Island, Philippines Bay of Campeche, Mexico Panglao Is., Bohol, Philippines ° These species a re proposed to be transferred from Lophiotoma to Unedogemmula (see text). purified using the PureLink PCR Product Purification Kit (Invitrogen Life Technologies, Carlsbad, California) following the manufacturer's suggested protocol. The eluted DNA fragments were digested with uracil specific excision reagent, annealed to pNEB206A vector (USER™ Friendly Cloning kit, New England BioLabs, Inc., Beverly, Massachusetts) and the resulting products tr wastorine d into « competent DEH5e cells ( Samnfrcle anc F.M. Heralde III et al., 2007 Russell, 2001). Plasmid DNAs were isolated from ampi- cillin resistant colonies and the nucleic acid sequences of the inserts determined using AB] DNA sequencer with ABI Big Dye chemistry (Foster City, DNA se- quences have been ie re to GenBank and the accession numbers are: EF467333, EF467333 EF467335, EF467336, EF467337, E splints EF467339, EF467340, EF467341, EF46734 EF467343, EF467344, EF467345. ae EF467347, EF467348, EF467349, EF467350, EF467351, EF467352, EF467353, EF467354, EF467355, and EF467356 ies >) fs ») 4 Sequence Analysis: — Nucleic acid sequences (the long- est of which had 593 nucleotides) were aligned manually using MEGA version 3.1 (Kumar, 2004). One tree was created from two independent runs using the software program MrBayes (Huelsenbeck, 2001: Ronquist, 2003). 5,000,000 trees were made in each run, 50,000 of which were saved. Two hundred and fifty of each of those 50,000 were also discarded as burn-in. Each run had four chains (one cold and three heated). The two independent runs were combined into a single tree where branches were preserved if they were found in 70% or more of those trees not discarded. The standard deviation after 5,000,000 generations was 2.401 x 10~°. A general time reversible (GTR) model was used, with the rate variation of some sites kept invariable and the remaining rates drawn from a gamma distribution. The other tree was created using the software program PHYML (Guindon, 2003). A thousand trees were obtained using non- parametric bootstrap analysis and combined into a single tree where branches were preserved if they were found in 70% or more of the given trees. A G TR model was used, with the base frequency estimates found empiri- cally and the proportion of invariable sites estimated. Four substitution rates were used, with the gamma dis- tribution parameter estimated. RESULTS AND DISCUSSION PCR Amplification of 12S Sequencing: The se- quences of 12S <7 from 23 species in the subfamily Turrinae (see Table 1) were obtained as described above. The 12S sequence of a Drillia species, Drillia regius Habe and Murakami, 1970), was used as the outgroup for the phylogenetic analysis. The sequences obtained are shown in Table 2: these were aligned for maximal overlap. Phylogenetic Analysis: — A phylogenetic tree, shown in Figure 1, was constructed as described under Meth- ods. The species that are presently assigned to two of the major turrine genera, Turris and Gemmula, appear as monophyletic clades in the phylogenetic tree obtained through Bayesian methods. However, the Lophiotoma species analyze d clearly split into two distinct, well- separated groups. Thus. the species presently assigned to Lophiotoma analyzed in this study do not appear to constitute a monophyletic assemblage. A large separation is found between two groups of Lophiotoma species; one group includes Lophiotoma unedo (Kiener, 1839 in 1834-80), Lophiotoma tayabasensis Olivera, 2004, Lophiotoma panglaoensis Olivera, 2004, Lophiotoma indica (Réding, 1798), and Lophiotoma bisaya Olivera, 2004. These spe- cies appear to be much more divergent from the Turris and Gemmula branches than the other group of Lophiotoma, which includes the type rie of Lophiotoma, Lophiotoma acuta (Perry, 1811); the latter comprises two cre branches, one branch pheal spe- cies such as Lophiotoma cingulifera (Lamarck, 1822), assigned by many syste matists to the sub yFenus Xenuro- turris Iredale, 1929, which is regarded as a separate ge- nus by some workers (Powell, 1966). Generic Classification and Nomenclature: = The un- expected phylogenetic separation between two groups of species conventionally assigned to the genus Lophiotoma, makes the present assignment of these spe- cies into the conventional Indo- Pacific turrine genera, Turris, Lophiotoma, and Gemmutla inconsistent with the phylogenetic tree shown in Figure 2. One potential so- lution would be to lump these Indo-Pacific genera to- gether under one genus, Turris, and use subgeneric des- ignations for each large clade of species (this might be called the “Conus alternative”; the major group of Indo- Pacific Turrinae comprise a phylogenetic branch that does not appear to be more dive ergent by molecular cri- teria than is the divergence within the species presently assigned to the genus Conus; Espiritu, 2001). ). Although this altermative may have some merit, the substantial lit- erature referring to species in the traditional genera Tur- ris, Gemmula, and nuns (including a significant paleontological component of the research literature) would make this a radical (and probably impractical) al- ternative. Because Lophiotoma acuta is the designated type spe- cies for the genus Lophiotoma, the species presently in Lophiotoma that are in the branch not including L. acuta require a new generic designation. There are two ge- neric/subgeneric designations potentially available for the group. One is a name proposed originally by Powell, Lophioturris Powell, 1964; Powell envisione d Lo} hiotur- ris as a genus allied to Lophiotoma with Loptanionis indica (Réding, 1798) as type. Lophioturris was set up specif Really { for forms that have blunt paucispiral proto- conchs; since most of the species in this clade have po- lygyrate (multispiral) protoconchs, Lophioturris does not seem to be an appropriate taxonomic designation (see Powell, 1964, for a discussion of differences in proto- conch morphology). The other available generic name for this group of species (which would have priority) is Unedogemmula MacNeil 1960: as originally proposed, Unedoge ~mmula was a separate genus, with l indogemmula une do (Kiener, 1839 in 1534-80) as type. However, P< well relegated Unedogemmula to be a subgenus of Gemmula. 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ODD. [€0Z] 3DD WLVOVLLVDD DLOVWYDODY SDLODOOLYO LOLOLYWOLL OWWOLYLOVY YWOLLVYLLL W-YLY-LYV- OVWLVLVLYL LLVVYWLOWWY WWOVLYLIOL LLIVS-LOL¥Y LLLLOWYOLL LYOLIY-¥-¥ LO---DYOVL LOVOVOYWYY WLOOLYYOWY -LYLYOLOYY LIYYWOVWD09 youLL99999 aL “sp. OA DYPAC ‘FG ‘pgp piyshodg “CG (OMMDSOL DINUUIE ) ‘GG juolosyy D]NUUE Mm) 1G DJNOIV DUO} doy ‘0G ‘Bsowads DjmuUay ‘GT ‘vapauowp DNUUAD “ET ‘s{suapodos YJNUAUAE) “LT “WSUOUID: 1OspK : CL “SIpupad smn] rl pruojle 04 SLLENT, °C] sipgqojoods stung “Sp syphych dot OT ia pwuojor fouojorydoq yp vdo.sphjod puojorydory -g ‘opoun puojporydoy °G ‘swsuaspqvhv, puojorydory “fp ‘sisuaosunjo muojorydory ‘e “paafynsuyo puojonydor is “sruiofinyee 10 puojordory ‘T ‘soouanbos WNC SZ] Jo JUSWUSYY "Zs ATQULh F. M. Heralde III et al., 2007 Page 135 xt BS % > = a £8 & cf > Ss = 5 2S < %, ae ae S “a, a i} —“ = < Aa 4 SS 3 gS eo “ee = = € of WY % “ BE 8 FF — % fF Sa 8k ¥ es cA ey 38 5 & Ra Phiotom, es Po 48 St S| Ch tiphylis gt? so ov gv” Lophiotoma polytropa : re oi? e i wo" emmula p OSario \ of ,sal aM Q 2 yw" we? § Ge wo FS aN NY 8 ( + iS ef og gz & 3 = v & SI Ny K s Ss SS g Figure 1. Phylogenetic tree of species listed in Table 1. A tree based on 12S rDNA sequences was constructed (see Methods) Species from three Indo-Pacific families in the subfamily Lei were analyzed (Turris, Gemmula, Lophiotoma). In addition, one Atlantic aes Polystira albida and a species in the subfamily Drillinae (or Clavinae), Drillia regius were included. Note that the species of Lophiotoma are split into two widely separated groups. As discussed in the text, species marked by an asterisk are proposed to be moved to another genus. Lophiotoma jickelii and Turris grandis are non-standard taxomomic assignments : the figured specimens: these are widely regarded as synonymous to Lophiotoma acuta and Turris crispa, respectively (see Powell, 1964). However, the molecular data clearly se parates Lophiotoma acuta and Lophiotoma jickelii, and the type of Turris crispa is sufficiently divergent from the form shown that using Turris grandis seems a preferable name for this species. quently, Kilburn (1983) suggested that based on general shell morphology, Unedogemmula seems much closer to species traditionally assigned to Lophiotoma, and that it should more appropriately be regarded as a subgenus of Lophiotoma; this suggestion has been adopted in most of the recent taxonomic treatments of turrine genera. The molecular results reported above demonstrate that Unedogemmula unedo is indeed closely related to some of the spe cies most taxonomists prese mitly include in Lophiotoma, (such as Lophiotoma indica, Lophiotoma bisaya. and Lophiotoma tayabasensis). Furthermore, the molecular analysis clearly shows that there is no justifi- cation for designating Unedoge mmula as a subgenus of either Gemmula or Lophiotoma, since Unedogemmula unedo is in a very divergent branch of the phyloge netic tree. Thus, our results support the original designation of Unedogemmula as a full genus, although the species comprising the genus need to be somewhat redefined. Unedogemmula unedo is the type species, and the larger, strongly maculated forms previously assigned to Lo} shiotote (such as Lophiotoma indica, Lophiotoma ee nsis, and Lophiotoma bisaya) are transferred to Unedogemmula from Lophiotoma. A recent analysis of Philippine forms related to these species, clarifying the relationships between these forms and the Unedogem- mula unedo group, was recently published (Olivera, THE NAUTILUS, Vol. 121, No. 3 ee eee ee ee ee Drillia regius Polystira albida oii i \o Unedogemmula panglaoensis aa 98 70 Unedogemmula unedo 84 95 98 96 91 100 Turris babylonia ah a Seca Unedogemmula tayabasensis Unedogemmula indica =X. Turris garnonsii Aa Turris grandis —-= Turris spectabilis al 100 Turris totiphyllis =: Gemmula diomedea = [we 100 86 Gemmula speciosa 83 100 85 400 100 a | 0.1 expected substitutions per site Moure 2 Figure 2. generations). Note that all the major groups are highly supp assigned to Lophiotoma is now regarded as a full genus, Unedogemmula, with Unedogemmula unedo as type figure, these species (st 2004 . ated in that work a nedogemmutla, as rede fine: d below, ae this should it seems likely ong all of the species level taxa » properly placed in the genus be veritied by obtaining molecular data for those species. Genus Unedogemmula MacNeil, 1960 30-LO5mm, fusiform with anterior canal unnotched Shell large straight, Description: tall s toconc h variable often with the transitional part of the larval shell deco- pire long Pro- from blunt paucispiral to multispiral, rated with brephic axials or axially costate whorls. Sinus ill labeled Lophiotoma in Figure 1) are Gemmula sogodensis ~~ ap Gemmula lisajoni ee NN Gemmutla rosario 100 The phylogenetic tree in Figure 1 is re-plotted, except that the confidence limits are included (calculated after 10 million wrted; as is explained in the discussion, one of the groups formerly species; thus, in this designated as Unedogemmula spp. is peripheral, deep, and narrow at the termination of the sinus rib. Remarks: ~The genus has shell morphology with strong similarities to Lophiotoma: most spt ae s have a smooth peripheral keel but, in some there are distinct peripheral granulations. In other species, these granula- tions continue to the body whorl, but these tend toward obsolescence in most forms. In contrast to Lophiotoma, the peripheral keel does not consist of two raised cords at in spe cle S, the edges with a depress sd area in the center; rather, F. M. Heralde III et al., 2007 Page 137 some forms, there is a single smooth rib exhibiting a variable level of peripheral granulation. In order to test the veracity of the proposals based on the Bavesian analysis and outlined above, a second ap- proi ach to the phy logenetic an: ysis of the sequences was also carried — A Maximum Likelihood method was employed as described under Me thods. The results of se analysis are shown in Figure 3. The separation of the species described above origin: ily in Lophiotoma into two distinct groups is strongly sup porte ed using this analy- . Thus, both methods support raising Unedoge mmula The analyses differ, Gemmula; the maximum . a full genus, as described above. however, in the results with likelihood method does not group all of the species ana- lyzed into a single monophyletic clade, but into two groups of species. Thus, given this discrepancy between the two methods, the monophyly of Gemmula clearly requires further investigation. A brief summary of a proposed revision of Indo-Pacific genera in the subfamily Turrinae is given in Table IIL. The cladogram in Figures 2 and 3 give support to Turris and Unedogemmula. However, Lophiotoma (redefined to exclude the species transferred to Unedogemmutla) has two branches, both strongly supported; the conjoining of the branches has less than 90% support in the Bayesian analysis, and is not supported above the cutoff level Drillia regius Polystira albida Unedogemmula panglaoensis Unedogemmula tayabasensis Unedogemmula unedo 97 72 85 99 87 100 Unedogemmula indica Unedogemmula bisaya 84 Turris normandavidsoni Turris garnonsii Turris grandis Turris babylonia Turris spectabilis Turris totiphyllis Gemmula diomedea 84 Gemmula sogodensis Gemmula speciosa 100 97 95 100 98 Figure 3. Phylogenetic tree of species listed in Table 1 the tree in Figure 1, but was constructed using the PHYML This tree was made using the same sequence a software Gemmula lisajoni Gemmula rosario Lophiotoma polytropa Lophiotoma jickelii 97 Lophiotoma acuta Lophiotoma olangoensis Lophiotoma cerithiformis Lophiotoma cingulifera lgnment as that used for program - > IAQ Page 135 THE NAUTILUS, Vol. 121, No. 3 (70%) in the Maximum Likelihood analysis. Given these data, the solution would be to split Lophiotoma into two separate genera, Lophiotoma and Xenuroturris. We feel that at this time the more conservative approach of re- taining the genus Lophiotoma, and dividing it into two subgenera, Lophiotoma (s.s.) and Xenuroturris (with Lophiotoma (Lophiotoma) acuta and Lophiotoma (Xe- nuroturris ) cingulife ra as types, respectively) is prefer- able until a wider range of species has been analyzed. There are a number of species (presently in Lophiotoma) that are problematic to assign (such as Lophiotoma ruth- veniana (Melvill, 1923)), and we believe that a molecular analysis of these forms needs to be carried out before we fully understand the relationship between Lophiotoma (s.s.) and Xenuroturris. It may well turn out that, when the eae is completed, the separation between the two branches (Lophiotoma and Xenuroturris) will be defini- tive; at that point, separating the two groups of species into different genera will be justified. The major conclusion from this work is that Unedogemmula should be recognized as a full genus, and is a sister group to the major branch that includes Turr is, Gemmula, and Lophiotoma (as redefined). ACKNOWLEDGMENTS We thank Sean Christensen for help with experiments Bradford Stevenson for help with the preparation of phy- logenetic trees and Drs. Edgar P. Heimer de la Cotera and Estuardo Lopez-Vera for providing the Polystira sample. This work was ea ed in part by a Program Project grant from the National Institutes of Healthy, GM48677. LITERATURE CITED Bouchet, P., P. Lozouet, P. Maestrati, and V. Héros. 2002. Assessing the magnitude of species richness in tropical marine environments: high numbers of molluscs at a New Caledonia site. Biological Journal of the Linnean Society 75: 421-436. Bouchet, P. and J. P. Rocroi, 2005. Classification and nomen- clature of gastropod families. Malacologia 47: 1-397. Bouchet, P., A. Sysoey, and P. Lozouet. 2004. An inordinate fondness for Turrids. In: Molluscan Megadiversity: Sea, Land, and Freshwater, [Abstracts of the] World Congress of Malacology, Perth, Abstracts, p. 12. ee D. J. D., M. Watkins, V. Dia-Monje, G. E. Cartier, L. J. Cruz, and B. M. Olivera. 2001. Venomous cone snails: molecular NU ae and generation of toxin diversity. Toxicon 39: 1S99-1916. Guindon, S. a O. Gascuel. 2003. A simple, fast and accurate algorithm to estimate large phylogenies by maximum like- lihood. Systematic Biology 52: 696-704 Huelsenbeck, J. P. and F. Ronquist. 2001. MRBAYES: Bayes- ian inference of phylogeny. Bioinformatics 17: 753-755. Kilburn, R. N. 1983. Turridae (Mollusca: Gastropoda) of south- ern Africa and Mozambique. Part 1. Subfamily Turrinae. oo of the Natal Museum 25: 549-585 —_ A. J. 1998. Superfamily Conoidea. In: Beesley, P. L., >. J. B. Ross, and A. Wells (eds.) Mollusca: The Southern nee Fauna of Australia. CSIRO Publishing, Mel- bourne, pp. 546-854, Kumar, S., K. Tamura, and M. Nei. 2004. MEGA3: Integrated Software for Molecular Evolutionary Genetics Analysis and Sequence Alignment. Briefings in Bioinformatics 5; 150-163. Olivera, B. M. 2004. Larger forms in Lophiotoma: Four new species described in the Philippines and three from else- where in the Indo-Pacific. Science Diliman 16; 1-28. Oliverio, M. and P. Mariottini. 2001. A molecular framework for the phylogeny of Coralliophila and related muricoids. Journal of Molluscan Studies 67: 215-224. Ponder, W. F. and A. Warén. 1988, Classification of Caenogas- tropoda and Heterostropha—A list of family-group names and higher taxa. Malacological Review, Suppl. 4: 288-328. Powell, A. W. B. 1964. The family Turridae in the Indo-Pacific. Part 1. The subfamily Turrinae. Indo-Pacific Mollusca 1: 297-346. Ronquist, F. and J. P. Huelsenbeck. 2003. MRBAYES 3: Bayes- ian phylogenetic inference under mixed models. Bisse matics 19: 1572-1574. Sambrook, J. and D. W. Russell. 2001. Molecular Cloning—A Laboratory Manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. Taylor, J. D., Y. I. Kantor, and A. V. Sysoev. 1993. Foregut anatomy, feeding mechanisms, relationships, and classifi- cation of the Conoidea(=Toxoglossa)(Gastropoda). Bulle- tin of the Natural History Museum, London (Zoology) 59: 125-170. Watkins, M., D. R. Hillyard, and B. M. Olivera. 2006. Genes expressed in a Turrid venom duct: divergence and simi- larity to conotoxins. Journal of Molecular Evolution 62: 247-256. THE NAUTILUS 121(3):159-145, 2007 Page 139 Reproductive biology of the nudibranch Doris fontainei VOrbigny, 1835 (Gastropoda: Opisthobranchia) from the M agellanic Region Le Co Claudia Muniain' Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” Avda. Angel Gallardo 470 C1405D]R, Buenos Aires ARGENTINA cmuniain@macn.gov.ar Valdivia, CHILE Carlos S. Gallardo Instituto de Zoologia E. Universidad Austral de Chile Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales “Bermardino Rivadavia” Avda. Angel Gallardo 470 C1405D]R, Buenos Aires, ARGENTINA and Departamento de Biodiversidad y Biologia Experimental Facultad de Ciencias Exactas y Naturales Universidad de Buenos Aires Buenos Aires, ARGENTINA F. Kilian ABSTRACT The present study describes the egg deposition, embryonic and larval development, and reproductive behavioral traits of the South American dorid nudibranch Doris fontainei dOrbigny, 1835, observed in laboratory under controlled conditions. Be- havior during copulation and spawning was recorded using a digital video camera. Copulation and spawning lasted 24 h and 22-93 h, re spectively. The spawned mass reached 1.80 m in length and was coiled in a counter-clockwise direction from the centre with one edge affixed to the substrate. The ribbon con- tained numerous small capsules, each having 2-4 eggs of about S6 ym in diameter. Intracapsular egg development lasted about 14 days at 14.5°C, culminating with the release of up to 1.25 million veliger larvae per egg mass. The new veliger larvae showed incipient development of the foot and a relatively small protoconch (ca. 158 jm), indicating they enter a planktotrophic phase. A comparison with other nudibranchs, and particularly with dorids, suggests that D. fontainei has an annual cycle whose egg mass fits a morphological pattern typical of the fam- ily. The characteristics of its encapsulated development show it is one of the most fecund species among those having this pattern, which also explains, in part, its geographical dispersion from Peru in the Pacific to northern Argentina in the Atlantic. Additional Keywords: Nudibranchia, reproductive behavior, multiple embryos, planktotrophic development INTRODUCTION An interesting characteristic of nudibranch gastropods is their capacity to produce ee and delicate egg masses, which can be obtained in laboratory through ae equate maintenance of adult individuals. The pattern of early development of these egg deposits is a basic trait in : : Author for correspondence the life histories of the different species. These patterns, which can be de termined through observations under controlled conditions, may also provide new elements to better typify and distinguish species whose taxonomic status is still debated. Hurst (1967) has grouped the egg deposits of opistho- branchs into three morphological types in accordance with the morphology of the ribbon, mode of attachment, and alignment of the capsules. However, Thompson (1967) defined three developmental larval strategies: planktotrophy, lecithotrophy, and direct dev elopment. Although there is general knowledge about egg depo- sition and definition of some dev elopme ntal parameters for different species worldwide, the priate biology of the rich and diverse nudibranch assemblages inhabit- ing the coasts of South America is still poorly known. Doris fontainei dOrbigny, 1835, is a widely distributed and common guiiscach from the South American coast. This sea slug inhabits the extensive littoral fringe of the SE Pacific (Peru and Chile), spanning the entire Magellanic region, to latitude 38°S in northern Argen- tina, following cold Antarctic currents to depths of 70 m (Muniain et al., 1991; Muniain, 1997; Schrédl, 1997a, b: 2000). As with other dorids occurring along this stretch of coastline, D. fontainei has been subjected to several taxo- nomic revisions, including a recent re-assignment based on detailed anatomical studies (Valdés and Muniain, 2002). An exploratory sampling for nudibranchs on the coast near Valdivia, Chile (40°S), allowed us to collect and maintain living specimens of D. fontainei in the labora- tory and observe their reproductive behavior, including copulation, spawning, some aspects of individuals behav- ior after completion of egg deposition, and plank- totrophic larval stage. Page 140 THE NAUTILUS, Vol. 121, No. 3 C. Muniain et al., 2007 Page 141 MATERIALS AND METHODS A total of six specimens of Doris fontainci were collected from Los Molinos (40°S), Valdivia, (Chile), by scuba div- ing to depths of 3-6 m a 4-16 March 2001. The water temperature at the sampling site was 14-15°C. To obtain information on the copulatory and reproductive behavior of Doris fontainei, the specimens were maintained in aquaria containing aet rated seawater at approximately 14.5°C, at the Zoological Institute of the Universidad Austral de Chile, Valdivia. Egg masses deposited on the aquarium walls were cultured in the original container without dislodging the egg mass. Routine isothermal seawater changes were made daily with water filtered to 1 um. Egg dev elopment was monitored until hatching. Egg samples were preserved in 6% formalin in seawater for further microscopic studies, which included counting and measuring the capsules and the eggs and embryos contained inside them. Most of the information pre- sented in this study was recorded by photography and digital video during maintenance of living individuals. Complementary bhsernations were made using pre- served material. Voucher material was deposited in the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” under accession number MACN 36542. Egg deposits were described from macroscopic obser- vations as well as using a Zeiss Axiostar stereoscopic mi- croscope equipped with a digital camera and imaging software at Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”. The number of eggs per capsule were determined using repeated counts in randomly chosen locations throughout of the egg ribbon. Develop- ment was observed under microscope and each develop- mental stage illustrated, particularly that of the troco- phore and middle vel iger; mean numbers of embryos per capsule, and their size relation to the size of the capsule were ca Iculated. Capsular volume (V) was determined (in wm”) based on measurements of capsular length and width, assuming that the depth axis was equiv alent to the width axis. The radii (r), ro, r3), and the volume was calculated using the formula: = 4/37 x 1 x Yo X Is. Sixty capsules previously separated in groups according to the number of embryos they contained were 1 randomly extracted from the gelatinous matrix and measured as mentioned above. The diameters of 40 eggs at the unsegmented zygote stage were measured. Descriptions of ihe dev elopment included three embryonic stages: trochophore, middle to advanced veliger, and pre-hatching veliger; each stage was photographed and measured (n = 30). The total number of eggs in a deposit was estimated indirectly by counting egg capsules in two different 5 mm-wide seg- ments of the egg ribbon. For this purpose representative segments of ihe ribbon were isolated by cutting them W ith a scissor and carefully extracting individual capsules from the gelatinous mi itrix of each segment. The mean number of eggs per cm of eg¢ ribbon was determined, and the total number of eggs in the deposit was calcu- lated based on the length of the entire ribbon. The rib- bon length was detesmiadd by measuring the length of its whatiached free edge from the photogr aphic images, taking into account the photo scale. The approximate number of embryos in the egg ribbon was estimated multiplying the number of egg capsules by the average number of embryos they éontained: the total number or embryos was estimated following the same procedure as for eggs. The development period was determined in an egg deposit maintained under controlled temperature, and was considered to start at egg deposition and finish at the initiation of hatching. Recently hatched larvae were observed under stereoscopic microscope. RESULTS Copulatory and Spawning Behavior: Copulation and post-copulation behavior of two of the six speci- mens collected and maintained in aquaria was observed (Figures 1-8). During copulation, nudibranchs had the rignt sides in contact at the level of the genital See with the heads in opposite directions (Figure 1). Copu- lation lasted between 2 and 4 hours, with the animals in total contact; penises were not visible because they were obscured by the mantle, although there was a conspicu- ous penial papilla, as described by Muniain et al. (1991) and Valdés and Muniain (2002). One individual 76.2 mm in length began to spawn in a counter-clockwise direction eraistial) ) at 16:00 h on 13 Mar. 2001 and fin- ished at 15:00 h on 14 Mar. 2001 (23 h) at a temperature of 14.7°C. During spawning, movement was very slow, with the rhinophores contracted within the sheaths and the branchial tuft exposed but motionless. Following deposition of the egg mass in the center of the aquarium, the nudibranch slowly moved to one side. A second specimen, 64 mm in length, began to spawn two hours after collection on 16 Mar. 2001, showing sinistral ovi- position, and terminated at 11:00 on a Mar. 2001, after a total spawning time of 22 h (Figure 2). Following com- plete spawning, the slug cemmained ss oa arebile: contracting and completely invaginating very slowly, with only the posterior portion of the foot adhered to the Figures 1-8. Doris fontainei 7 Orbigny, 1835, reproductive behaviour captured from digital video. 1. Copulation maintained in aquaria. Scale bar = 1 cm. 2-3. Specimens showing sinistral oviposition (counter-clockwise direction). Scale bars: Figure 2 = 1 cm; Figure 3 = 1.5 cm. 4. Detail of the post-spawned behavior. Scale bar = 1 cm. 5. After half and hour immobile, the nudibranch slow] returning to a normal position. Scale bar = 0.5 mm. 6. Detail of rhinophores and gills exposed again. Scale bar = 1 cm. 7. Detail of the free border and scattered spaces of the spiral ribbon. Scale bar = ri, rhinophores; fo, foot em. Abbreviations: gi, gills: ma, mantle: mo, mouth: ri 2 cm. 8. Complete spawn, specimen (64 mm). Scale bar = 1.5 ‘ > 9 Page 142 THE NAUTILUS, Vol. 121, No. 3 ” lll * Figures 9-13. Doris fontainei @ Orbigny, 1835, 9-12. Details of the ovoidal capsule with 2 and 3 eggs. The volume of the capsule increases with the number of embryos, 3 embryos is the most common number. Scale bar = 45 jam. 13. Hatching planktotrophic larva, right lateral view. Typical coiled shell (type 1, Thompson, 1961). Scale bar = 25 pm. operculum; rm, retractor muscle; se, statocyst; v, velum. substrate (Figures 3, 4). The ventral surface of the mantle comple tely covered the organism, which raised so as to show the anterior portion of the foot. The nudi- branch remained in this position for about half an hour (Figure 5), slowly returning to a normal position, extend- ing the rhinophores, cinnalline the mantle, and slowly crawling away from the egg deposit ( (Figure 6). This be- havior was completely re corde d with digit: u video from 11:00 to 12:10 hrs on 17 Mar. 2001 (Figure 8). Egg Mass and Larval Development: The egg mass formed a spiral ribbon of concentric rings consisting of five to seven revolutions around a central point (Figures The free border of the ribbon was undulated, un- like the attached border, which was straight (Figure 7). The undulations of the free border coincided with con- centric rings of the ribbon. Both the egg deposits ob- tained in the laboratory and those collected in the field were pink. The latter were typically positioned under rocks or in shaded areas. The egg mass ae some mucous areas without egg capsules (Figure 7). The egg capsules were arranged line arly along the el abn. and were joined by a fine cord (chalaza) that served to maintain the spiral configu- ration. Randomly scattered spaces without capsules oc- curred in the egg ribbon and in its different median portions. The egg de posit from the 64-mm specimen was formed in six revolutions around the center, with a maxi- mum diameter of 10 cm; it was 13.4 mm in height, and an estimated 1.8 m in length (Figure 8). Mean egg diameter was 86.5 + 4.7 xm. The capsules contained multiple embryos, although single embryos appeared occasionally The number of e smbryos in cap- sules ranged from 2 to 5,3 e ee . ing ‘clearly the most common number ( Figures 9— The first count based on 338 capsules, containing fe ee gave an average number of 3 embryos per capsule. A second count, of 361 capsules (containing intermediate veligers), 1300 Vv al Abbreviations: int, intestine; op, produced an average number of 3.18 embryos per cap- sule. The average of the 2 counts was about 3.1 embryos per capsule. The a tended to be ovoid, exhibiting some more irregular shapes when crowded. The volume of an egg capsule increased with the number of embryos it contained (Figures 9, 11). As expected, the eggs demon- strated typical spiral cleavage. Measurements of the different stages were as follows: trochophore = 99. 35, - 5.5 jum (n = 20), advanced encapsulated veliger = 116.25 + 125 55 pm (n = 20), and hatched veliger larva = 158.1 + 7.16 wim (n = 10). Hatching of typically planktotrophic veligers becha 14 days after incubation, at an avet rage temperature of 14. 5°C. These veligers measured a mean of 150 jzm (n = 20) in prostomial length, with the pro- toconch sinistrally coiled, with 4 to 1 whorl at hatching. At hatching, the larvae had no eyes nor propodium ru- diment, although the velum, velar retractor muscle, lar- val kidney, nephrocy sts, operculum, and cephalopedal alimentary apparatus were developed (Figure 13), The total number of larvae released from the egg mass was estimated indirectly through observations under controlled conditions. Microscope- aided counts carried out on 2 transects of the egg ribbon showed an average of 2236 egg capsules per cm. With an estimated length of ca. 1.8 m (indicated above), the entire egg deposit con- tained about 402450 capsules, and with the above esti- mate of 3.1 embryos per cé ipsule, the entire egg deposit contained about 1.25 million embryos, resulting in the same number of developed planktotr yphic veliger larvae - a few days later. DISCUSSION The present study describes some of the traits that typify the the history of D. fontainci, specifically in regard to its reproductive pattern. Its relatively large body size sug- C. Muniain et al., 2007 Page 143 gested a priori that the species would have an annual evcle, with one generation per year, and the spawning characteristics fell into Class A, ‘following the classifica- tion and corresponding attributes of dorid nudibranchs prov ided by Hurst (1967) The characteristics of the e neapsulated development described in the present study agree with the embryonic developmental pattern observed in most of the Dorida- cea, based on information summarized by Thompson (1967) that typifies the three basic developmental stages observed in that group of nudibranchs. The embryonic development observed in D. fontainei fits Type-1 pattern of this scheme. Free planktotrophic larvae are released at hatching, there are multiple relatively small embryos per capsule, and the period of e smbryonic development lead- ing to hatching of weakly dev eloped veliger larvae is comparatively short. Based on information on related species (Thompson, 1958: 1967: Strathmann, 1987: Goddard, 2005) most dorids with planktonic life cycles have eggs ranging be- tveen 60 and 130 jm in size, many of which are in the 70-90 zm range, in agreement with values found for D. fontainei. There is also agreement in the number of em- brvos produced per spawning, with values as high as several hundred thousand embryos per spawning in the most fecund species, such as Acanthodoris pilosa ( coe gaard, 1789) and Archidoris pseudoargus (Rapp, 1 gh (both reviewed by Thompson. 1967). peer fontaine is a highly fecund dorid, given that it ee over a million eggs per spawning. This notably exceeds the numbers reported for other species in the group; the closest species is A. pseudoargus, which spawns 645000 eggs (Colgan, 1914, cited in TI soot a 1967). The du- ration of embryonic development (pre-hatching) in planktotrophic species can be as short as 6 to 10 days, and rarely exceeds one month (Thompson, 1967). The development period of 15 days found for D. fontaine: falls within the range expected for dorids with plank- totrophic dev elopment. Another trait coinciding with the Type-1 pattern of development are the characteristics and degree of development of the veliger larvae hatched from the egg deposits of D. fontainei. As Thompson (1967) and Todd et al. (2001) mentioned for species showing this pattern, typical characteristics include ab- sence of eyes or propodial rudiment, but presence of a deve sloped velum, retractor muscle, larval kidney, oper- culum, and cephalopedal digestive system, which were the most notable traits observed by us. Based on observations of lecithotrophic dorids of the genera Adalaria, Dendronotus, Discodoris, Tritonia (Thompson, 1958; 1961; 1962; 1976; Gohar and Abul-Ela, 1959: Roginsky, 1962; Todd, 1979) and of the direct-developing species Cadlina laevis Linnaeus, 1767 Roginsky, 1962: Thompson, 1967), the eggs are 200 and 400 j.m in size, respectively, and the total number of embryos per spawn is clearly lower (aprox. 15000 to 50000), reaching only a few ‘hundreds in species with direct development. As indicated above, it is clear that D. fontainei has multiple embryos per egg capsule, but given the lack of information, we do not iow if this condition is common within Doridacea, although Thompson (1967) mentions it as a frequent characteristic among the opisthobranchs that have planktotrophic larvae and Type-1 develop- ment. Among the studies of reproductive aspects in other species of M: agellanic opisthobranchs, the presence of multiple embryos was noted in the saccoglossan Elysia patagonia Muniain and Ortea, 1997, which completes development with the formation of planktotrophic larvae (Muniain and Ortea, 1997; Muniain and Penchaszadeh, 2000; Muniain et al., 2001). Given the large number of embryos that D. fontainei produces, eadlonine of more than one embryo per egg capsule is probably an efficient mechanism for reducing the energetic cost of these egg masses compared to the relatively large benefits of maximizing the numbers of larvae gener rated per spawning, Although there is little information available on some species in this region, the development of D. fontainei can be compared with that of other dorids that oceur on the Chilean coast, such as Peltodoris marmorata (Bergh, 1898) (cited as Anisodoris rudberghi). It has eggs of SO wm in diameter and a larger number of eggs per capsule (5-23), as well as a pre- hatching veliger of 125 jum in length, and its embryonic development occurs in only 10 days at 14-16°C (Brokordt, 1992). The large number of larvae produced by D. fontainei suggests a possibly high degree of larval mortality and, aesouiated with this, a potentially effective mechanism for extensive larval dispersal. Wide dispersal can main- tain gene flow among populations as well as the extensive geographical dispersal (Schrédl, 1997a, b; 2000; Valdes and Muniain, 2002) shown by this nudibranch in com- parison with other species on the South American coast. There are no records in the literature of detailed ob- servations on the behavioral mechanisms accompanying spawning of nudibranchs. The intriguing post-hatching behavior shown by a specimen of D. fontainei in the present study deserves further laboratory research with a higher number of individuals to determine whether be- havior has a given pattern at this stage of the reproduc- tive process. The duration of the spawning process (22-23 h) asso- ciated with the relatively long, highly fecund egg ribbon of D. fontainei may imply the generation of some facts of phy siological imbalance, whieh the animal strives to over- come through the behavioral event observed at the cul- mination of oviposition. Whether this behavior results from an inordinate energy demand compared to that occurring in other species of nudibranchs is unknown. Evaluations made in prosobranchs have shown he high relative energetic cost involved in making benthic egg masses (Perron, 1981; 1982). Studies on the chemical ecology of this species have shown that it biosynthesizes repellent compounds that accumulate only in the mantle tubercles (Muniain, 1997 Page 144 THE NAUTILUS, Vol. 121, No. 3 Gavagnin et al., 1999). If this behavior is confirmed un- der laboratory conditions, further studies should also evaluate its possible occurrence in natural conditions, where the nudibranch would be exposed to predation throughout the entire stage. Nudibranchs can be considered semelparous (despite spawning repeatedly in a season) in that once they have reached maturity, their period of spawning is inevit ably followed by apparent genetically programmed post- reproductive e death (Todd et al., 2001). There fore, repro- duction must be fully successful in the single opportunity presented to these animals, which will probably not be able to survive until another reproductive season to re- peat the process. Such is the high cost of reproductive activity for the individual. The bvoad geographic distri- bution of D. fontainei makes it a useful specie s for future studies of pc yssible geographic variations in its pattern of embryonic development. Further studies including re- lated nudibranch species in the region should be con- ducted. For example, studies of the dorid Adalaria proxima (Alder and Hancock, 1854) have shown that the egg diameter and other embryonic and larval traits could show intraspecific variation, reflecting the adjustment of its populations to a given range of variability in environ- mental conditions where the species is distributed (Jones et al., 1996; Todd et al., 2001). ACKNOWLEDGMENTS This study was funded by the GEF-BIRF 28385/AR (A- CB-51), CONICET (PIP 5301), ANPCyT (PICT 34111) and DID-UACH (S2005-12) ) proje cts. C.M. and P.E.P are Research Members of the National Research Council of Argentina (CONICET). LITERATURE CITED Brokordt, Kk. 1992. Intensidad reproductiva y desarrollo intra- eee uw de tres especies de nudibranquios presentes en la IV Region de Chile (Anisodoris rudberghi, Phidiana inca y Thecacera darwini), y su relaci6n con la estrategia de alimentacion larval. Thesis, Universidad Catélica del Norte Coquimbo, Chile. Gavagnin, M., N. Uneur, F. ¢ G. Cimino. 1999. New minor diterpenoid diacylglycerols Jastelluecio, C. Muniain, and ae the skin of the nudibranch Anisodoris fontaini Jour- nal of Natural Products 62: 269-274. Goddard, J. H. 2005. Developmental mode in benthic opistho- branch molluscs from the northeast Pacific Ocean: feeding in a sea of plenty. Canadian Journal of Zoology 82: 1954— 968 Gohar, H. A. F. and I. A. Aboul-Ela. 1959. On the biology and development of three nudibranchs (from the Red Sea). ublication of the Marine Biological Station Al-Ghardaqa 0: 41-62 Hurst, A. 1967. The egg masses and veligers o thirty northeast acific opisthobranchs. The Veliger 9: 255-288 Jones, H. L., C. D. Todd, and W. J. Lambert. 1996. Intraspe- cific variation in embryonic and larval traits of the dorid nudibranch molluse Adalaria proxima (Alder & Hancock) around the northern British Isles. Journal of Experimental sb eB iOwey and Ecology 202; 29-47, Muniain, C., J. Ortea and G. Rodriguez. 1991. Redescripcién de Ne oa is carvi Marcus, 1955 de las costas de Patagonia, con notas sobre las relaciones entre los géneros Arc hidoris y Neodoris. Iberus 10: 105-111. Muniain, C. 1997. Moluscos Opistobranquios de Argentina: Revision Taxonémica y relacion de Ecologia Quimica en algunas especies patagonicas. PhD Thesis. University of Oviedo, Spain, Muniain, C. and J. Ortea. 1997. First record of a sacoglossan (=ascoglossan, Opisthobranchia) ) from Patagonia (Argen- tina); Description of a new species of genus Elysia Risso, 1818. Veliger 40: 29-37. Muniain, C. and P. E. Penchaszadeh, 2000, Development pat- tern in the sacoglossan Elysia patagonica Muniain and Ortea, 1997 from Argentina. Abstracts of 66th Meeting of the American Malacological Society and 33rd Meeting of the Annual Western Society of Nalacolosicts San Fran- cisco, p. SI, Muniain, ©., A. Mari, and P. E. Penchaszadeh. 2001. Ultra- structure of the digestive gland of larval and adult stages of the sacoglossan Elysia patagonica, Marine Biology 139; 687-695. Perron, F. E. 1951. The partitioning of reproductive energy between ova and protective capsule s in marine ¢ gastropods of the genus Conus. American Naturalist 11S: 110-118. Perron, F. E. 1952. Inter and intraspecific patterns of repro- ductive effort in four species of cone shells (Conus spp.). Marine Biology 68: 161-167. Roginsky, I. S. 1962. The egg-masses of nudibranchs of the White Sea. Biology of the White Sea 1: 201-214. Schrédl, M. 1997a. Range extensions of Magellanic Nudi- branchs (Opisthobranchia) into the Peruvian faunal prov- ince. The Veliger 40; 35-42. Schrédl, M. 1997b, On the morphology of the Magellanic nudi- branch Anisodoris fontaini (D’Orbigny, 1837), and its syn- onymy with A. tessellata Bergh, 1898. The Veliger 40: 228-233. Schrédl, M. 2000. Taxonomic revision of the common South American nudibranch Anisodoris fontaini (D°Orbigny, 1837), with discussion of its systematic placement. Journal of Molluscan Studies 66; 49-61. Strathmann, M. F. 1987. Reproduction and Development of Marine Invertebrates of Northern Pacific Coast. Univer- sity of Washington Press, Seattle, pp. 268-302, Thompson, T. E. 1958. The natural history, embryology, larval biology and post-larval is velopment ‘of Adalaria proxima (Alder and Hancock) (Gastropoda, Opisthobranchia). Philosophical ae of the Royal Society of London (Series iB Biological Sciences) 242; 1-58. Thompson, T. E. 1961. The importance of the larval shell in the classification of the Sacoglossa and the Acoela (Gas- tropoda, ( ypisthobranchia). 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Valdés, A. and C. Muniain. 2002. Revision and taxonomic re- assessment of Magellanic species assigned to Anisodoris Bergh, 1898 (Nudibranchia: Doridoidea). Journal of Mol- lusean Studies 68: 345-351, THE NAUTILUS 121(3):146-158, 2007 Page 146 Two new modern records of the southern oyster drill Stramonita haemastoma floridana (Conrad, 1837) in Chesapeake Bay, USA Juliana M. Harding! Department of Fisheries Science Virginia Institute of Marine Science Gloucester Point VA, 23062 USA jharding@vims.edu M. G. Harasewych Departinent of Invertebrate Zoology, MRC-163 National Museum of Natural History Smithsonian Institution, Washington, DC 20013- Harasewych@si.edu P.O. Box 37012, 7012 USA ABSTRACT Live southern oyster drills, Stramonita haemastoma floridana (Conrad, 1837), have been collected from two Chesapeake Bay western shore tributaries. Four specimens were collected be- tween Brown Shoal and Thomas Rock in the lower James River in February 2005. Thirteen live southern oyster drills were collected from Back River in April 2006, Identification of these drills as Stramonita haemastoma floridana has been confirmed using DNA bar-coding data. Southern oyster drills collected in Chesapeake Bay are genetically nearly identical to populations from the Atlantic coast, and differ signific: uitly from popula- tions from the Azores and from the Gulf of Mexico. These collections mark the first reported records of live southern oys- ter drills from within the Chesapeake Bay estuary. It is un- known if these drills represent isolated introductions or expan- sions of the northern range this species. Water temperature patterns in Chesapeake Bay and the Mid-Atlantic Bight from 1990-2005 are similar to those observed in the late 1950s when Stramonita haemastoma floridana was first found living in Chincoteague Bay, Maryland. Additional Keywords: Range extension, water temperature, Cape Hatteras, zoogeography, DNA bar-coding INTRODUCTION The southern oyster drill Stramonita haemastoma flori- dana (Conrad, 1537) is a predatory gastropod typically found in association with populi itions of the oyster Cras- sostrea virginica Gmelin, 1791 (e.g., Burkenroad, 1931; sutler, 1955), A population from the Atlantic coast of the United States was described as Purpura floridana Con- rad, 1837, but later regarded to be a subspecies of the Author for correspondence broadly ranging Stramonita haemastoma (Linnaeus, 1767), a morphologically diverse taxon that has been re- ported to span the temperate and tropical coasts of the eastern and western Atlantic as well the eastern Pacific. The name Stramonita haemastoma hacmastoma (type locality: Tenerife, Canary Islands, fixed by Clench, 1947: 76) has been applied to the populations ranging from the Channel Islands southward to Senegal, the western Mediterranean, the Madeira, Canary and Cape Verde Islands in the eastern Atlantic (Clench, 1947: 76; Poppe and Goto, 1991: 141), the Azores (Morton et al., 1998), Trinidad southward to Uruguay in the western Atlantic (Clench, 1947: 76), and from Baja California southward to Peru in the eastern Pacific by some (e.g., Clench, 1947) but not all (e.g., Keen, 1971: 549) researchers. Stramonita hae Se floridana (type locality: Hy- poluxo Island, Lantana, Florida, see Clench, 1947: 77) was reported to range from North Carolina southward to Yucatan and the West Indies, and throughout the Ca- ribbean as far south as Trinidad (Clench, 1947: 76), extending to Brazil (Abbott, 1974: 180), The name oe monita hae smastoma canaliculata (Gri Ly, 1839) (type lo- cality originally but erroneously listed as China) has been applie 1d to a distinctive morphological variant prevalent throughout the Gulf of Mexico (e.g., Abbott, 1974: 180). Some three dozen names have been variously partitioned and synonymized for the many geogr iphically circum- scribed shell phe ce be longing to the Stramonita hacimastoma complex (see Rose abe ra, 2005). North American records of Stramonita have generally been divided into two subspecies, Stramonita haema- stoma floridana from the Atlantic coastline, and Stra- monita haemastoma canaliculata from the Gulf of Mexico; such division is based on shell characters (Ab- bott, 1974: Butler, 1985). Stramonita haemastoma canaliculata has been differentiated morphologically on J. M. Harding and M. G. Harasewych, 2007 Page 147 Figures 1-9. Stramonita spp. 1-6. Stramonita haemastoma floridana (Conrad, 1837). 1. USNM 618840, Willis Wharf, Hog Island Bay, Northumberland Co., Virginia, trapped in 2.4 m depth. September 15, 1955, ex-F. W. Sieling. 2. USNM 1091020, oyster reefs just below Deep Creek, Lower James River, Virginia, in approximately 3 m, February 2005. 3. Male dnd, female specimen, USNM 1091021. Back River, Virginia, in commercial crab pots set in 3 m, April 2006. 5. USNM 416874, Fort Macon, North Carolina. 6. USNM 1099258, Sand bar on north side of Fort Pierce Inlet, E of Little Jim Island, Ft. Pierce, Florida. 6 August 2004. 7. Stramonita canaliculata (Gray, 1839), USNM 1099259, St. Andrews Bay, Florida, rip-rap near campsites. 8. Stramonita haemastoma haysae (Clench, 1927), USNM 568834, Barataria Bay, Louisiana. 9. Stramonita haemastoma haemastoma (Linnaeus, 1767), USNM 1099260, intertidal rocks, Vila Franca do Campo, Sao Miguel, Azores, 26 July 2006. Page 148 THE NAUTILUS, Vol. 121, No. 3 Atlantic Ocean Atlantic Ocean Figure 10. Map of the Atlantic Ocean depicting the southeastern coast of the United States, Chesapeake Bay region [inset], and the Azores Islands, showing collection sites for Stramonita specimens and water-temperature data. Sites are identified as follows: Delaware lightship and buoy stations (DE), Chincoteague, VA (A1), Chesapeake lightship and buoy stations (C), Fort Pierce Inlet, FL (A2), St. Andrew's Bay, FL (A3), Pensacola, FL (A4), and Sao Miguel, Azores (A5). Within the Chesapeake Bay region Chincoteague, VA (B1), Maryland DNR monitoring station NBM1S49 (B2), Maryland DNR monitoring station NBM 1301 (B3), Virginia Institute of Marine Science, Gloucester Point, VA (B4), Back River, VA (B5), James River near Deep Creek (B6). the basis of its larger size, presence of strong, rugose shoulder nodules, and its deeply channeled suture. Sev- eral authors (e.g., Butler, 1954; Gunter, 1979: Walker, 1982) had reported that differences in shell morphology are neither consistent nor concordant with geographical patterns and concluded that subspecific distinction was not warranted. More recently Liu et al. (1991) con- firmed that characters of shell and radula were not taxo- nomically informative, but demonstrated that Stramonita from the northern Gulf of Mexico could be differentiated from populations of the Atlantic coast at “a level that is characteristic of congeneric species” using allozyme elec- trophoresis. Other recent studies have explored the ef fectiveness of the east Florida ecotone as a barrier to gene flow between Atlantic and Gulf coast populations for a variety of taxa (for reviews, see Avise, 2000; Wise et al., 2004). Vermeij (2001; 701) reviewed the Recent and fossil species of Stramonita, and suggested that the Stra- J. M. Harding and M. G. Harasewych, 2007 Table 1. Samples of Stramonita spp. used in this study. Taxon Locality Page 149 Voucher Number of (1) Pensacola, FL St. Andrew's Bay, F Sao Miguel, ees Stramonita canaliculata Stramonita canaliculata Stramonita haemastoma haemastoma Stramonita haemastoma floridana Stramonita haemastoma floridana Stramonita haemastoma floridana Deep Creek, VA Back River, VA Ft. Pierce Inlet, FL specimens specimens GenBank USNM sequenced accession number SSST09 1 US6330 1099259 1 EU073061 1099260 2 EU073051—EU073052 1099258 2 EU73053-EU073054 1091020 3 EU073055—E U073057 1091021 3 EU07305S—E U073060 " Sequence data from Harasewych et al., 1997 monita haemastoma complex may consist of as many as four western Atlantic and two eastern Atlantic species. He also noted similarities (bifid crenulations along the outer lip) between some specimens of S. canaliculata and S. bifida Vermeij, 2001, from the Cantaure Formation (early Miocene, Venezuela). Although the northern limit of Stramonita along the eastern United States has generally been reported as Or- egon Inlet, North C selina (Wells and Grey, 1960; But- ler, 1985), living individuals (Figure 1) and recent his- cee specimens (empty shells) were collected from lo- cations in the Maryland and Virginia waters of Chincoteague Bay (Figure 10, Al) in 1955 and 1956 (Sieling, 1955; 1960), extending the northern range boundary for this animal along the US Atlantic coast. Sieling (1960) suggested that the southern 6yster drills were ceded. into Chincoteague Bay with trans- planted oysters. Subsequent surveys of Assate: ague Island and Chincoteague Bay by Counts and Bashore (1991), Table 2. Homer et al. (1997), and Prezant et al. (2002) reported living S. haemastoma floridana from these habitats as febenily as 1996 (Prezant et al., 2002). Counts and Bashore (1991) categorized S. haemastoma floridana as “rare” a linked the relative decline in abundance and distribution of this drill with the decline in local oyster resources from 1960 to 1989. Prezant et al. (2002) report living S. haemastoma floridana in their survey and col- lected specimen(s ) from Memorial Park, Maryland. Living species ’ Stramonita have not previously been reported from the interior of the Chesapeake Bay (An- drews, 1956; Wass, 1972) in modern time, although Sh. canaliculatum had occurred in Chesapeake Bay, Mary- land, and S. h. floridana had reached southern New Jer- sey during the exceptionally warm Sangamonian Stage (80, 000 to ca. 220,000 yr BP) of the Pleicracene | Petuch 1997: 53). Ruiz et al. (2000) ) reported S. haemastoma as an eceblehed aes in the Chesapeake Bay region on the basis of the Chincoteague Bay, Maryland- Virginia, Summary of water temperature data sources from 1950-2005 for Delaware (DE, Figure 10) and Chesapeake (C, Figure 10). Sea surface temperature (SST), bottom temperature (BT) and air temperature at the water surface (AT) were used. Station Year(s) Data Station type Depth Source Delaware/Winter Quarter 1955-60 SST, BT Winter Quarter lightship 24-29 m ] 1961-70 SST, BT Delaware lightship 30 m 2 1970-S4 SST One degree quadrangles NA 3, 4,5 Chesapeake 1984-2005 AT, SST NOAA buoy 44009 28 m 6 1950-197] SST Chesapeake lightship 20 m 7,8 1971-84 SST One degree quandrangles NA 3, 4,5 1985-2005 AT, SST NOAA buoy CHLV2 11.6m 9 1. Winter Quater lightship data archive. East Coast USCG lightship/Lightstations. MBLWHOI Library data archives, Woods Hole, MA. http://dlaweb.whoi.edw/lightship/lightships_winterqtr.html 2. Delaware lightship data archive. East Coast USCG Lightship/Lightstations. MBLWHOI Library data archives, Woods Hole, MA. http://dlaweb.whoi.edu/lightship/lightships_delaware_html 3. 1966-1974. The Gulf Stream. U.S. Naval Oceanographic Office, Vols 1-9. 4. 1975-1980. gulfstream. U.S. Dept. of Commerce, NOAA, National Weather Service. Vols. 1-6. 5. 1981-94. Oceanographic Monthly Summary. U.S. Dept. of Commerce, NOAA, National Weather Service/National Earth Satellite Service. Vols. 1-14. 6. Delaware Bay, Buoy 44009. http:/Avww.nodc.noaa.gov/BUOY/44009. html. 7. Bumpus, D. 1957. Surface water temperatures along Atlantic and Gulf coasts of the United States. U.S. Fish and Wildlife Service Special Scientific Report—Fisheries No. 214. 5. Chesapeake lightship data archive. MA. http://dlaweb.whoi.edu/lightship/lightships_chespstn.html East Coast USCG Lightship/Lightstations. MBLWHOI Library data archives, Woods Hole 9. Chesapeake Light. VA, Buoy CHLV2. http:/Avww.node.noaa.gov/BUOY/chlv2. html. Page 150 THE NAUTILUS, Vol. 121, No. 3 SS SS SS eS eee [ 2005: James River, VA WB 2006: Back River, VA 9b ee 0 } lI [tet 64 "66 68 70 72 74 76 78 80 "82 "ga Ps 88 "90 a ea Midpoint of shell length class (mm) Number of snails collected J Figure 11. Length frequency distribution of live Stramonita haemastoma floridana specimens collected from Back River, VA (2006) and the James River, VA near Deep Creek (2005). and Hog Island Bay, Virginia, collections reported by Sieling (1960) (P. Forowolk Smithsonian Environmental Research Center, personal communication). Distribution patterns of many molluscan species along the US Atlantic coast are directly related to water temperature with Cape Hatteras acting as a natural zoogeographic boundary (Franz and Mervill, 1950a, b). Wells and Gray ( (1960) reported S. h. floridana on two subtidal shipwrecks south of Cape Hatteras, North Carolina, but found none on the four shipwrecks they examined north of Cape Hatteras. Wells (1961) described re soular collections of resident S. h. floridana from inte srtidal oyster reefs in the vicinity of Beaufort and Cape Lookout, North Carolina. Southern oyster drills begin laying eggs at water tem- peratures between 20 and 30°C (Stickle, 1999). At water temperatures less than 10-12.5°C southern oyster drills stop feeding on bivalves, bury into the substrate, and become inactive (Bulter, 1954; Gunter, 1979; Garton and Stickle, 1950; Stickle, 1999) until water temperatures rises. While the upper lethal thermal limit for southern oyster drills is 35 to 40° C (Brown and Stickle, 2002), the lower lethal thermal limit for southern oyster drills is unknown. This report documents the first collections of living Stramonita haemastoma floridana from within the Chesapeake Bay estuary. We characterize both oe of southern oyster drills collected in the Che ssapeake Bay genetically (cytochrome c oxidase I “DNA bar-coding”) aad compare ‘these data to se quences derived from rep- resentative spe cimens from the southeastern (Indian River Inlet) and northwestern (St. Andrew's Bay, Pensa- cola) coasts of Florida, as well as from samples from Sao Miguel, Azores. The presence of these drills in Chesa- peake estuarine habitats is analyzed in the context of a long term (1950-2005) coastal ‘bottom water tempera- ture data set and two independently collected estuarine bottom water temperature data sets (Chincoteague Bay, Maryland and Virginia, 1951-56; McGary and Seiling, 1953; Seiling, 1957; York River, Virginia, 1986-2005: VIMS, 2006) in order to relate the observed Chesapeake collections with environmental/climate conditions in known drill habitats along the Atlantic coast and in Chin- coteague Bay. MATERIALS AND METHODS Sample Collection: Four living S. haemastoma (Fig- ure 2) were collected in the lower James River, Virginia, in the Che ssape rake Bay from oyster (Crassosteea vir ginica) reefs just below Deep Creek (Figure LO, BG) at a depth of approximately 3m in Fe bruary 2005 and were tured in to the Virginia Institute of i Science (VIMS) rapa whelk [Rapana venosa (Valenciennes, 1846)| bounty program (see Harding a Mann, 2005, for bounty program details). All four Stramonita speci- Table 3. Nucleotide (above diagonal) and amino acid (below diagonal) ) ditfe srences between samples in the portion of the cytochrome c oxidase I gene obsoleta {Genbank NC 007781]}) (591 aligned positions, corresponding to positions 73 to 664 in the complete mitochondrial genome of Iyanassa Sample l 2 3 5 6 7 8 9 10 lI 12 1) S. canaliculata Pensacola, FL — 0) 126 126 123 123 123 122 124 124 123 122 2) S. canaliculata St. Andrew’s, FL 0) — 126 126 123 123 123 122 124 124 123 122 3) Sh. h aemastoma Azores | s S — 5 56 55 55 58 56 56 56 56 1) S. h. haemastoma Azores 2 i 7 | — 57 56 56 61 57 57 DT 57 5) S. h. floridana Ft. Pierce Inlet, FLI 7 7 3 2 ~ 2, 2 7 3 5 3 3 6) S. h. floridana Ft. Pierce Inlet, FL2 7 if 3 2 0) 0 5 | 3 | | 7) S. h. floridana Deep Creek, VAI 7 7 3 2 0 0 — 5 | 5 | | 5) S.h. floridana Deep Creek, VA2 S S 2 3 | | | —- 6 S 6 6 9) Sh. floridana Deep Creek, VA3 7 4 3 2 0 0 0 | = { 2 2 10) S. hh. floridana Black River, VAI v4 7 3 2 0 0 0) | 0 — { | 11) S.h. floridana Black River, VA2 7 7 3 yy) 0 0 () | 0) 0 = 2 12) S. h. floridana Black River, VA3 7 7 3 2 0) 0) 0 | 8) 0 0 J. M. Harding and M. G. Harasewych, 2007 Page 15] : licul S. canaliculata PenFL Gulf of Mexico S. canaliculata SAnFL S. haemastoma haemastoma Azores 1 Azores S. haemastoma haemastoma Azores 2 S. haemastoma Fort floridana FPIFL1 Pierce S. haemastoma Inlet, FL floridana FPIFL2 S. haemastoma floridana DCVA1 Deep S. haemastoma Creek floridana DCVA2 VA , S. haemastoma floridana DCVA3 S. haemastoma floridana BRVA1 Back S. haemastoma River floridana BRVA2 VA S. haemastoma floridana BRVA3 Figure 12. Strict consensus of four most parsimonious trees (1 = 176; ci = 0.955; ri = 0.960) resulting from an exhaustive search using maximum parsimony (PAUP 4.0b10) based on 591 bp of cytochrome c oxidase I sequences. Branch lengths from one of the four most parsimonious trees are above the branches, bootstrap (bold) and jackknife (bold italic) propor- tions are below the branches. mens were collected in a single oyster dredge tow. Thir- teen living S. haemastoma (Figures 3-4) were collected in Back River, Virginia, (Figure 10, B5) in April 2006 and were also turned into VIMS via the rapa whelk bounty program. The 2006 collection was made using commer- cial crab pots (wire mesh cubes with approximately 0 0.5m sides) baited with northern quahogs |Mercenaria merce- naria (Linnaeus, 1758)| deployed at a depth of 3 m. Shell lengths of specimens (mm, maximum dimension tip of the spire to bottom of the siphonal canal) were measured upon receipt at VIMS. Voucher specimens from both Chesapeake collections have been deposited in the Na- tional Museum of Natural History. Smithsonian Institu- tion (USNM, Table 1) ; DNA Bar-coding: Three specimens from each Chesa- peake collection were frozen and transported to the labo- ratory, where DNA was extracted from red buccal muscles of each animal using Qiagen DNEasy kits ac- cording to the manufacturer's protocol. A portion of the mitochondrial gene for the DNA bar-coding region of the cytochrome c oxidase I gene was amplified using Sigma JumpStart eee Re sadly Mix and Meyer's (2003) degenerate sd versions of the Folmer et al. (1994) HCO and LCO primers for the samples listed in Table 1. The resulting PCR products were cleaned using magnetic beads [Agencourt, manufacturer's protocol] and se- quencing reactions were run on ABI 3730 sequencers that were set up according to manufacturer's instruc- tions. Sequences were manually checked/corrected and assembled using Sequencer™ 4] (Gene Codes C Jorp.) then aligned wih Clustal W (Higgins et al., 1994) using default settings, and their pace analyzed using PAUP 4.0b10 (Sivoftord. 2002). Pairwise comparisons of sequences and amino acids were performed using Mega 3.1 (Kumar et al., 2004). Water Temperature Data: CoastaL HaAsirats: Wa- ter temperature data from 1950-2005 were used to de- scribe mean monthly bottom water temperatures (BT, C°) for two stations in the Mid-Atlantic Bight (Delaware/ Winter Quarter and Chesapeake) (Figure 10, Table 2). Daily sea surface and bottom water temperature data from the Delaware/Winter Quarter (1961-70) and Chesapeake (1958-71) lightship stations were used to calculate the average monthly observed difference be- tween sea surface tempe rature (SST) and bottom water temperature (BT) on a site-specific basis. The observed average monthly SST-BT differences from the daily lightship data were used to estimate average monthly B’ r at these sites during years after 1971 when only SST was available (Table 2). ne air temperature (AT) and SST data from NOAA buoys (1985-2005) were used to calculate the observed dteronoe between average monthly AT and average monthly SST for Delaware Light and Chesa- peake I Light. The observed av erage aioe AT-SST dif- ferences from the buoy data were use sd to estimate hourly SST for these sites during months after 1954 when only AT was available. Average residuals for annual BT from the long-term (1950-2005) average annual BT were calculated for each site in which at least 9 months of data were available. Monthly average bottom water temperatures (with stan- dard error of the mean) were calculated for each of 12 months for all sites. Monthly BT estimates from 1970 to 1983 (Delaware) and 1984 (Chesapeake Light.) use a single published monthly average ( Table 2) while monthly BT estimates from NOAA buoys (Table 2) are averages calculated from hourly readings with n values >400 per month. Estuarine Habitats: Water temperature data col- lected at Public Landing, Maryland, in Chincoteague Bay from McGary and Sieling (1953) and Sieling (1957) were used to calculate average monthly water tempera- tures (with standard error of thé mean) during the period 1951-1956. Modern (2001-2004) water temperature data from Chincoteague Bay stations south of Robins Marsh (XBMS8149) (Figure 10, B2) and near the MD-VA border (XBM1301) (Figure 10, B3) were obtained from Page 152 THE NAUTILUS, Vol. 121, No. 3 A. Delaware Feb 1961 - Dec 1970 102, 138170 148 1 2 3 4 5 B. Chesapeake Jan 1958 - Dec 1971 Average monthly SST-BT difference (°C) n= 202 188 174 180 169 1 2 3 4 6) eae e 13. 162-164 7 8 9g 10 11 12 ) e 145 197 159 173 7 8 i) 10 11 12 Month ae rage monthly difference between sea surface temperature (SST) and bottom water temperature (BT) from lightship data (Table 2) for Delaware (A) and Chesapeake (B) lightships. Error bars indicate standard error of the mean. N values (number of sail, SST-BT pairs used to estimate the SST-BT differences) at each site are presented above the X axis in each panel. the Maryland De Lain nt of Natural Resources Eyes on the Bay web site (http://mddnr.chesape: akebay. net/ bay_cond/). Water de pth at all (historical and modern) of these Chincoteague Bay sites is 3 m or less. Seiling (1954) reported less than a degree Celsius variation be- tween surface and bottom water temperatures and the data presented in McGary and Seiling (1953) and Seiling (1957) from. sites throughout the estuary confirm the well-mixed nature of these non-channel habitats. The Virginia Institute of Marine Science (VIMS) at Gloucester Point, Virginia (Figure 10, B4) maintained a water temperature monitoring station from 1986 through September 18, 2003, (arrival of Hurricane Isabel) that recorded bottom water temperatures (°C) ata depth of 2-3 m. The VIMS Molluscan Ecology program has main- tained an environmental monitoring station located within 200 m of the original VIMS station since January 2005 that records bottom water temperatures (°C) at depths of 2 m. Hourly water temperature averages were obtained from the VIMS data archive (1986 to Se p 2003) and the VIMS Molluscan Ecology program environmen- tal data archive (2005) and used to calculate monthly residuals from the 1986-2005 mean and average monthly bottom water temperatures (with standard error of the mean). RESULTS aa ad The southern oyster drill specimens col- lected from the James River in Febru: wy 2005 ranged in size from 66.4 to 75.6 mm shell leneth with an average shell length of 69.0 + standard error of the mean 2.9] mm. Sointhe rm oyster drills collected in April 2006 from Back River had shell le neths ranging from 73 to S9 mm with an average shell length of $3.3 + 1.41 mm. Size frequency distributions for both collections ( Figure 11) indicate that all specimens were adults (sexually mature; Burkenroad, 1931; Butler, 1985) and represent the up- per end of the size distribution typically found in Loui- siana habitats (Brown and Richardson, 1987; Brown et al,, 2004). These large individuals have probably reached a size refuge from most local predators including blue crabs (€ allincetes sapidus, see the work of Turra et al., 2005, using C. eve preying on S. hacmastoma) and are certainly capable of cating oysters >50 mm shell height (Garton, 1986; Brown and Richardson, 1987). J. M. Harding and M. G. Harasewych, 2007 A. Delaware Jan 1984 - Dec 2005 n= 10261 10050 12342 11677 12880 12493 13543 14049 12911 13168 11876 11666 2 3 4 5 B. Chesapeake Jan 1985 - Dec 2005 ) Average monthly SST-AT difference (C) 7 8 ) 10 11 12 n= 10317 9982 11059 10995 11258 12175 12422 11965 11671 11434 12023 11430 1 2 3 4 5 Figure 14. (Table 2) for Delaware (A) and Ches: apeake ( 7 8 9 10 11 12 Month Average monthly difference be Ae en sea surface temperature (SST) and air temperature (AT ) from NOAA buoys 3) buoys. Error bars indicate standard error of the mean. N values (number of daily SST-AT pairs used to estimate the SST-AT he at each site are presented above the X axis in each panel. DNA Bar-coding: Comparisons of a 591 base-pair portion of the auitechotd! il gene for cytochrome c oxi- dase I from the samples listed in Table 1 revealed that specimens of Stramonita collected within Ches sapeake Bay were genetically very similar or identical to speci- mens colle ecte 2d from Fort Pierce Inlet, on the southeast- ern coast of Florida, but differed substantially from Azorean samples and even more so from specimens from the northwestern coast of Florida (Table 3). Maximum parsimony analyses of the 144 phylogenetically informa- tive sites using the exhaustive search algorithm yielded four equally parsimonious trees (length = 176; ci = 0.955; ri = 0.960). Figure 12 illustrates strict consensus tree of these four trees, including results of bootstrap and jack- knife analyses. Water Temperature Data: =CoastTaL Hapirats: Aver- age annual bottom water temperatures in the period 1950-2005 for the two stations along the US Atlantic coast followed a latitudinal trend and were lower 9.81°C. SEM = 0.15, 600 months of data) at Delaware than at Chesapeake (12.30°C, SEM = 0.21, 596 months of data). The differences between SST and BT recorded by the lightships (approximately 1956-71) show the sea- sonal deve thom nt of the thermocline at Delaware and Chesapeake beginning in April and persisting until Oc- tober with the most pronounced differences between surface and bottom water temperatures occurring in July and August (Figure 13) when these stations experience surface temperatures that are at least 7° C higher than bottom temperatures. Air temperatures re eee d by = Delaware and Chesapeake NOAA buoys (T: able 2 are 1-2° warmer than SST from April through July (Fig. ure 14). Examination of annual average residual bottom water temperatures from the average long-term (1950-2005) bottom water temperatures at coastal stations (Figure 15) shows that both sites see above average wa- ter temperatures during the late 1950s. Multiple con- secutive years between 1970-1980 and, again, in the pe- riod 1995-2005. Estimated bottom water te mperatures for 2002 were among the highest observed during the period 1950-2002 for Delaware and Chesapeake Bays Delaware and ¢ Jhesapeake, both north of ¢ vape Hatteras, experience water temperatures of 10°C or less eight and five months out of the year, respectively (Table 4A Page 154 THE NAUTILUS, Vol. 121, No. 3 —_ 2 A. Delaware oO —_ 3 = © i 8 2 € oO ~ 5 2 1 ° co} oO © 5 O > © wo S aI -1 j=) wo (o>) = = -2 je) = § 3 (3 n oO on TOO ON TOWONTSOWDWDIONTGOAOGOA TLC OM]M OA +t ag ouonwnonwnonwnooeDnoogoaorRrenRRRRWWAADANAD DD DAD OS OF DADDADAADA AAD AAA ADAAAADAAAAAAADAADAADAA DAA OO AS Sar oe el er ool, so ei, a Ton, a Se, od, Seal Sen a a ee eR co on oe, ec ed Te) | ~ B. Chesapeake ae P ~~ Oo — = = 2 o 2 Qa = o = E = i P= fe) co} D > 0 — oO > © wo So -1 oO Py fo) o 2 2 = (eo) = Go - g -3 Ke) oO oNWNnNWONMONWOODADAARe RE RERR DWWADWDADWANANBDDDDHD OO OS w DOOD ADADAADA ADA AA ADA A DAA AAAAAADAAAAIAAADAA HA OOO TS SS YS wwe SSS SS SS SE SS SS SS SS SS SE SES ON NN Year Figure 15. Annual average residual bottom temperature (BT) from the long term (1950-2005) average bottom temperatures for Delaware (A) and Chesapeake (B). Long term average data for each site are presented in Table 4. The error bars represent the tandard error ol the mean mM degrees ¢ elsius J. M. Harding and M. G. Harasewych, 2007 Table 4. Summary of average monthly bottom water temperatures for coastal stations (A, 1950-2005) and estuarine stations (B) shown in Figure 10 and discussed in text. Standard error of the mean for each value is given in parentheses. A. Month Delaware Chesapeake Jan > as (0,22) 6.96 (0.24) Feb 5 (0.20) 5.60 (0.23) Mar 5 58 (0.18) 6.12 (0.21) Apr 7.02 (0.15) 8.31 (0.20) May bora 29) 11.02 (0.21) Jun 3 (0.18) 14.34 (0.21) Jul 9 45 (0.15) 14.93 (0.20) Aug 10.69 (0.20) 16.72 (0.25) Sep 14.62 (0.29) 20.27 (0.20) Oct 16.56 (0.20) 19.11 (0.21) Nov 13.57 (0,15) 11.02 (0.21) Dec 10.06 (0.18) 10.41 (0.23) B. Chincoteague Bay, Chincoteague Bay, Chincoteague Bay, York River, Public Landing, MD XBMS8149 XBM1301 Gloucester Point, VA Month (1951-56) (2001-04) (2001-04) (1956-2005) Jan 4.14 (0.36) 5.3 5.7 5.32 (0.36) Feb 6.22 (0.33) 5.25 5.38 5.55 (0.47) Mar 9.76 (0.35) 9.65 99 8.46 (0.33) Apr 14.57 (0.43) 15.53 15.85 13.61 (0.30) May 20.1 (0.38) 18.65 18.87 18.86 (0.26) Jun 24.08 (0.38) 2120 26.33 23.91 (0.26) Jul 27.77 (0.32) 26.1 25.85 26.96 (0.18) Aug 27.2 (0.26) 27.42 27.65 27.11 (0.18) Sep 23.6 (0.40) 22.95 22.93 23.99 (0,59) Oct 17.32 (0.50) 14.55 14.6 19.28 (0.33) Nov 10.72 (0.38) 15.1 15.68 13.56 (0.24) Dec 5.64 (0.39) 5.2 5.5 $8.36 (0.54) EsTUARINE Hapirats: Annual average residual bottom water temperatures from the York River at Gloucester Point, Virginia were higher than the 1986-2005 annual average (16.23 + 0.56°C) in 10 of the 1S years for which data are available (Figure 16) including 2005. The relatively shallow (3 m or less) estuarine habitats examined in Chincoteague and Chesapeake Bays expe- rience a seasonal average monthly water temperature cycle appropriate for their latitudes (Table 4B) with wa- ter temperatures less than 10°C observed December through March. The same pattern was observed in Chin- coteague Bay during 1951-1956 (Table 4B). DISCUSSION The collections of live Stramonita from the James River (2005) and Back River (2006) mark the first modern record of this predatory gastropod from the Chesapeake Bay interior. The morphology of these specimens, par- ticularly their larger size, deeply channeled suture, and the presence of strong, rugose shoulder nodules in the Back River samples, is similar to that of some populations from the northern Gulf of Mexico [particularly the form named Stramonita haemastoma haysae (Clench, 1927 (Figure 8)|. This, in tum, has raised questions about whether these animals were introduced into Chesapeake Bay, and the possible source of such introductions. How- ever, the results of the “DNA bar-coding” study clearly uicaeate that the Chesapeake Stramonita haemastoma floridana are genetically very similar to, and in one case indistinguishable from, a population of Stramonita hae- mastoma floridana trom southeastern Florida. There seems little doubt that the source of the Chesapeake Stramonita haemastoma floridana populations lies along the eastern seaboard of the United States. Less clear is whether these populations were introduced into Chesa- peake Bay with oysters, as had been suggested by Sieling (1960) for the Chincoteague Stramonita, or if their pres- ence is due to a northward expansion of the range of Stramonita haemastoma floridana due to warmer ocean temperatures. The “bar-coding” data also indicate that the east-coast Stramonita are well differentiated from, yet more similar to Stramonita from the Azores than to specimens from the northern Gulf of Mexico (Figure 12), Provisional taxonomic consequences of this study are to recognize the Azorean populations as Stramonita haemastoma hae- mastoma, to retain the usage of Stramonita haemastoma Page 156 THE NAUTILUS, Vol. 121, No. 3 3 York River, Gloucester Point, VA -2 Residual from 1986-2005 average bottom temperature (C) (jo) 1950 1952 1954 1956 1958 1960 1962 1964 1966 1968 1970 1972 1974 Figure 16. Annual average residual bottom temperature (16.23 ° of the mean in degrees Celsius. floridana for the populations from the eastern United States, and to distinguish the populations from the north- erm Gulf of Mexico as Stramonita canaliculata, cated by Liu et al. (1991). Detailed analyses of this spe- cies complex over its entire ge ographic range are clearly required to better unde rstand its bioge ography and tax- as ac lve )- onomy. Water temperature patterns in Chesapeake Bay habi- tats and along the coast of the lower Mid-Atlantic Bight the late 1990s have average as were water temperature trends observed in the period 1955— 1957 when live Stramonita were collected on the Atlantic coast of Maryland and Virginia (Sieling, 1960). Although the lower thermal lethal limit for southe ‘mm oyster Sills is the om water temperatures in Chincoteague Bay during the 1951-56 are within 1 to 2° Fibs ape aa observed in the lower York River from 1986 ave seasonal te mperature regime »5 conduc ‘ive to survivi al since been above unknown, fact that observed seasonal trends in bot- yeriod C of bottom water 2005 sugge sts that Che 'sapei ake Bay tributaries may Living southern oyster drills were col- Bay by Prezant et al. (2002) of this animal ected from Chincoteague during surveys between 1991 and 1996, some 40 years after the initial collection (Sieling, 1956). The very low sopulation levels (“rare”, Counts and Bashore, 1991) of Bay in local L991) rather than southern oyster drills observed 19SS—89 have ovster re in’ Chincoteague been attributed to a decline Ince sources (¢ ounts and Bashore owmoonrwtowdoondriewrTraoewWdeiondrst KER WAWDWDAWDAADADABDWDIWAABDHA GOO DAA AADAAADAA DAA HD OO OO bl ee ee el ee el ee ee ee ee oe Year (BT) from the long term (1986-2005) average bottom temperatures C, standard error of the mean 0.56 °C) recorded at Gloucester Point, Virginia. The error bars represent the standard error unfavorable water temperatures. The tributaries of the lower Ches: ape rake Bi My have resident populations of bi- valves commonly eaten by Stramonita including mussels and oysters ( (Butler, 1985: Garton, 1986: Brown and Ri- chardson, 1987). Several of these tributaries are also sites of focused oyster restoration efforts that incorporate ad- dition of either spat on shell or broodstock oysters fur- ther expanding the potential prey field for southern oys- ter drills. If these collections represent the beginning of an in- vasion into Chesapeake Bay tributaries, the persistence of Stramonita haemastoma floridana in Chesapeake Bay will be dictated by the rmal tolerances as subjected to seasonal temperature cycles, while the geographic distri- bution will be set by si linity tolerances of adult and lar- vae. Adult Stramonita haemastoma may survive at salini- ties as low as 5-7 ppt (Gunter, 1979; Stickle, 1999). Stra- monita haemastoma egg capsules survive and release viable larvae at salinities down to at least 7.5 ppt and possibly as low as 3.5 ppt (Stic ‘kle, 1999). Veliger larvae may survive up to 5 days when exposed to LO ppt (Wells, 1961) but Roller and Stickle (1989) reported very low survival at salinities less than 15 ppt. In June 1972, heavy rains from Tropical Storm Agnes in combination with summer temperatures killed essentially all of the native oyster drills [Urosalpinx cinerea (Say, 1822), Eupleura caudata (Say, 1$22)| in the upper re saches of Ches: apeake Bay tributaries and reset the distributional range of the J. M. Harding and M. G. Harasewych, 2007 Page 157 native drills to the high salinity, lower reaches of the Chesapeake Bay where natural oyster populations cur- rently persist in only limited regions and numbers. Re- establishment of these native gastropod species to their former habitats is occurring slowly over decadal time scales and is limited by the Trek of planktonic larvae for both Urosalpinx cinerea and Eupleura caudata. Reinva- sion of their historic habitats by the native drills is con- founded by the fact that the historically widespread dis- tribution of oyster reef habitat has been drastically re- duced in areal cov erage since the early 1960s by a combination of diseases and environmental degr: adation. Competition for the niche vacated by the native drills during Tropical Storm Agnes already includes one large non-native gastropod, the veined rapa whelk (Rapana venosa) (Harding and Mann, 1999; 2005), which has planktonic veliger larvae like Stramonita haemastoma and is equally long- lived. Regardless of how these Stra- monita haemastoma got to these C hesapeake tributaries, the presence of acide: individuals in these tributaries adds yet another competitor for this vacant niche as well as additional predation pressure on local oyster resources. Successtul invasion of a habitat requires a breeding population and the presence of all life history stages in the new habitat (Williamson, 1996). Southern oyster drills live from 5-20 yr in Florida (Butler, 1985) ‘with generation times on the order of 12 months (Burken- road, 1931; Butler, 1954). The small number of speci- mens collected to date combined with the absence of these animals in annual fishery independent surveys of oyster reefs in the James River conducted by Harding and colleagues at VIMS and the Virginia Marine Re- sources Commission since the early 1990s may be an indicator that the observed southern oyster drills speci- mens represent small, isolated introductions that have not yet established local populations. ACKNOWLEDGMENTS Special thanks are extended to all local watermen who have donated Stramonita haemastoma and Rapana venosa to the VIMS research programs. Karen Caposella, Christina Conrath, Meredith Fagan, Adriana Picariello, and Matt Robinson assisted with local whelk collections by VIMS. Dr. Gregory Herbert, University of South Florida, kindly provided specimens of Stramonita from St. Andrew’s Bay, Florida. Azorean specimens were col- lected during a workshop in Vila Franca do Campo, Sao Miguel, Azores, hosted by The University of the Azores. The Workshop was a joint organization of Sociedade Afonso Chaves and the Department of Biology of the University of the Azores. Support from FLAD (Portu- guese- American Foundation for Dev elopment) is grate- fully acknowledged. Specimens from the Indian River Inlet were collected during a workshop supported by the Smithsonian Marine Station at Fort Pierce. Dr. Katherine Farnsworth, Indiana University of Pennsylvania, provided GIS assistance with maps. This is Contribution Number 2859 from the Virginia Institute of Marine Science, Glouc- ester Point, Virginia, and Smithsonian Marine Station at Fort Pierce Contribution Number 696. LITERATURE CITED Abbott, R. T. 1974. American Seashells, 2"" edition. 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THE NAUTILUS 121(3):159-162, 2007 Page 159 Brachycythara beatriceae, a new species from the Alboran Sea and the eastern Atlantic Ocean (Gastropoda: Neogastropoda: Conidae) Paolo Mariottini Dipartimento di Biologia Universita di “Roma Tre” Viale Marconi 446, 00146 Roma, ITALY mariotpa@uniromas3.it ABSTRACT Based on shell characters, Brachycythara beatriceae, a new gastropod species of the family Conidae from the Alboran Sea and the eastern Atlantic Ocean, is here described. The new taxon, represented by five specimens dredged along the Span- ish Mediterranean and the West Sahara coasts, is compared with the similar Brachycythara atlantidea (Knudsen, 1952), a species that occurs in the same geographical area. This new species is conchologically distinct and can be identified on the basis of its teleoconch shape, rib count and microsculpture, protoconch morphology, and shell color. Additional Keywords: Mangeliinae, turrid, shell morphology. INTRODUCTION Traditionally, the epithet “turrid” has been used as a general term referring to the numerous species belong- ing to the family-group Turridae H. Adams and A. Ad- ams, 1853, sensu lato. The new taxon described in the article is assigned to the family Conidae Fleming, 1822, subfamily Mangeliinae Fischer, 1883, genus Brachy- cythara W oodring, 1928, in accordance with the revision of the super rfamily Conoidea ( = Toxoglossa) Rafinesque, 1815, as proposed by Taylor, Kantor, and Sysoey (1993). The genus was re-described by Powell ( 1966: 117), and consists of small Recent and Tertiary species that aa 7 occur in the Caribbean area. Brachycythara has been represented in the eastern Atlantic Ocean by only one species, Brachycythara atlantidea (Knudsen, 1952) (see Roldan and Otero-Schmitt, 1999), which has recently been reported from the Alboran Sea by Smriglio et al. 2007). I had the opportunity to examine five shells of a species that, in spite of their similarity with B. atlantidea, could not be considered that species. These shells, col- lected along the southern Spanish (Costa del Sol) and West Sahara coasts, show features typical of the genus Brachycythara. After a comparison with the similar and sympatric B. atlantidea, I believe they represent an un- described species. Abbreviations used are: MZB, Laboratorio di Malaco- logia, Museo di Zoologia dell Universita di Bologna, Italy. Private collections cited in this article are: CS-PM, Carlo Smriglio and Paolo Mariottini (Rome, Italy); SR Stefano Rufini (Rome, Italy); FG, Franco Gubbioli (Marbella, Spain). SEM photographs were carried out at the LIME (Interdepartmental Laboratory of Electron Microscopy), University “Roma Tre”, Rome, Italy. SYSTEMATICS Superfamily Conoidea Ratinesque, 1815 Family Conidae Fleming, 1822 Subfamily Mangeliinae Fischer, 1883 Genus Brac hycythara Woodring, 1928 Type Species: — By original designation, Cythara gibba Guppy, 1896. Woodring, 1925, Carnegie Institute Washington of Publications 385, p. 175. Brachycythara beatriceae new species (Figures 1-9; 13-20) Description: Shell rather small, to 10.9 mm in length, holotype 8.6 x 3.4 mm, biconical, elongate-fusiform, solid, spire about half total height. Protoconch multispi- ral, dome shaped, of 3-3.5 strongly convex whorls, first 1.5-2 whorls smooth, remainder wheds reticulated with oblique axial costae crossed by spiral ribs of about equal width; protoconch indicative of planktotrophic larval de- velopment. Diameter of protoconch about 700-750 jum. Protoconch-teleoconch transition not well marked. Te- leoconch of 5-6 whorls, weakly angulate near middle of spire, sutural ramp gently concave, whorl sides gently convex; last whorl about 2/3 shell length. Axial sculpture consisting of 7-S prominent, slightly opisthocline, flexu- ous, and narrowly rounded axial folds: folds regularly spaced, with much broader inte rspaces, ret aching from suture to suture on spire but fading out after crossing the Page 160 THE NAUTILUS, Vol. 121, No. 3 Figures 1-12. Shells of Brachycythara species. 1-3. Brachycythara beatriceae new species, holotype, 5.6 « 3.4 mm, MZB 31032, Spain, Alboran Sea, off Marbella, 36°25’ N, 4°52’ W, 30-40 m. 4-6. Brachycythara beatriceae new species, paratype A, 10.9 x 4.5 mm, CS-PM, Spain, Alboran Sea, off Malaga, 36°33’ N, 4°22' W, 50 m, 7-9. Brachycythara beatriceae new species, paratype B, 9.3 3.7 mm, FG, coast of West Sahara, 50-60 m, 10-12. Brachycythara atlantidea (Knudsen, 1952), specimen L, 9.5 x 3.6 mm, FG, coast of West Sahara, 30-60 m. Scale bars: 2 mm shouldes slope and at about the middle of the base. Spira sculpture of numerous, very fine threads that densely alternate with bigger interspaces; the subsutural threa shows well-marked axial denticles. At higher magnifica- tion, it can be observed that each interspace consists Oo several up to tive) row of rounded tiny granules, each one linked axially to the uppel and lower thread by a very Ine connection \pe rture narrow, ovate, about one thir of the shell height Siphonal canal short, narrow, anc opel Inne lip vith a moderately developed pariecta | | Outer lip thin or variced according to the we of growth, whether the lip coine ides with an axia 1 1 interspace Anal sinus marked, arcuate on ope. Shell color white vith a brown band on lower half of body whorl (juvenile and subadult shells uniformly white); a darker spot present on the outer lip at the boundary of the white and the brown colors Type Material: Holotype (Figures 1-3), 8.6 « 3.4 min, MZB 31032, Spain, off Marbella, 36°28’ N 4°52’ W, 30-40 m; Paratype A (Figures 4-6), 10.9 x 4.5 mim, CS- PM, Spain, off Malaga, 36°33" N 4°22" W, 30-40 m; Paratype B, 8.3 x 3.1 mm, SR, Spain, off Malaga, 36°33' N 4°22" W, 30-40 m; Paratype C (Figures 7-9; 13-20) 9.3 «3.7 mm, FG, West Sahara, 50-60 m: Paratype D 8.2 «x 2.6 mm, FG, West Sahara, 50-60 m Other Material Examined: Shells of Brachycythara atlantidea (Knudsen, 1952): Specimen A, 10.2 « 4.1 mm P. Mariottini, 2007 Page 16] te wed Roe Figures 13-28. Shell morphological details of Brachycythara species by SEM photographs. 13-14. Brachycythara beatriceae new species, paratype B. 15-20. Details of the larval whorls and the shell sculpture. 21-22. Brachycythara atlantidea (Knudsen, 1952 spec. F. 23-28. Details of the larval whorls and the shell sculpture Specimen B, 9.2 7 mm; Specimen C, 7.5 x 3.4 mm mens A—G, N are deposited in CS-PM collection, speci- Specimen D, $8.2 x 3.6 mm; Specimens A—D from Spain, mens H—M in FG collection off Estepona, 36°25’ N, 05°09" W, 150-250 m; Specimen E. 7.7 x 3.4 mm, from Spain, off Adra, 36°45’ N, 03°01’ W. S0-150 m; Specimen F (Figures 21-28), 7.4 x 3.2 mm: Specimen G, 10.2 x 4.4 mm; Specimens F—G from Distribution: Alboran Sea (Costa del Sol Spain and Spain, ott Malaga, 36°33’ N, 04°22’ W,50 m: Specimen eastern Atlantic Ocean (West Sahara H, 10.5 x 4.3 mm; I, 9.3 x 3.6 mm: Specimens H—I from Spain. off Marbella. 36°28’ N, 04°52’ W, 30-40 m Specimen L (Figures 10-12), 9.5 x 3.6 mm; M, 8.2 x 3.4 mm: Specimen N, 6.3 x 2.6 mm; Specimens L-N Etymology: This species is named afte: the author's dredged along the coast of West Sahara: 30-60 m Speci- daughter Ww OO Type Locality: Alboran Sea, Spain, off Marbella 36°28' N, 4°52’ W, 30-40 m depth Habitat: The dredged specimens were from muddy bottoms Page 162 THE NAUTILUS, Vol. 121, No. 3 Remarks: Brachycythara beatriceae new species shows several shell diagnostic features (shape and sculp- ture of the protoconch and the teleoconch) that match the ones described by Powell (1966: 117, plate 18, fig. 7) for the genus Brachycythara, an amphiatlantic group of Recent to Miocene/Pliocene species whose distribution ranges from the Caribbean to the coast of West Africa (Powell, 1966; Rolan and Otero-Schmitt, 1999). Brachy- cythara atlantidea (Figures 10-12, 21-28) has been the only species ae to this genus known to occur in the eastern Atlantic Ocean (Rolén and Otero-Schmitt, 1999) and in the Alboran Sea (Smriglio et al., 2007). Brachycythara beatriceae and B. atlantidea show close similarities, but the two taxa can be easily separated on the basis of several shell morphological differences, as summarized in Table 1. Furthermore, at high magnifi- cation the complicated spiral microsculpture of B. bea- triceae shows finer threads, smaller and more numerous granules in each interspace. The finding of three speci- mens of B. beatriceae from two localities off the Spanish coast (Alboran Sea) indicates that this species can be added to the Recent Mediterranean malacofauna, while the collecting depth suggests that it is a circumlittoral species. As a final conclusion, the genus Brachycythara is at the present represented by two sympatric species dis- tributed in the Alboran Sea and the eastern Atlantic Table 1. Comparisons of shell features between B. beatriceae and B. atlantidea B. atlantidea Morphological characters B. beatriceae Prontoconch diameter size 700-750 600-650 (in jzm) Protoconch number of whorls 3-34 3 Teleoconch axial folds 7-8 9-10 (last whorl) Shell color white with a basal brown uniformly yellowish- band brown Ocean, Brachycythara atlantidea and Brachycythara beatriceae. ACKNOWLEDGMENTS I am grateful to Mr. Carlo Smriglio and Dr. Marco Ol- iverio (Dipartimento di Biologia Animale e dell Uomo, Universita di Roma “La Sapienza”, Rome, Italy) for their critical comments and advice on the present paper. I would like to express my deep gratitude to Mr. Franco Gubbioli (Marbella, Malaga, $ Spain) and to Dr. Stefano Rufini (Dipartimento di Biologia, Universita di Roma “Tor Vergata’, Rome, Italy) for kindly supplying type material of B. beatriceae and specimens of B. atlantidea. Dr. Andrea Di Giulio (Dipartimento di Biologia, Univer- sita di “Roma Tre”, Rome, Italy) is acknowledged for SEM photographs, which were carried out at the LIME (Interdepartmental Laboratory of ie Microscopy, Universita di “Roma Tre”, Rome, Italy). I am very in- debted to two anonymous referees for Shr suggestions and corrections to the manuscript. LITERATURE CITED Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and Museum 5: 1-157. Rolan, E. and J. Otero-Schmitt 1999. The F roa Turridae s.. (Mollusca, Neogastropoda) in Angola, 2. Subfamily Man- geliinae Fischer, 1883. Argonauta 13: 5-26. Smriglio, C., A. Di Giulio, F. Gubbioli, and P. Mariottini 2007, Brachycythara atlantidea (Knudsen, 1952) (Gastropoda, Neogastropoda, Conidae), first report from the Western Mediterranean Sea. Basteria, 71(1—3): 1-4. Taylor, J. D., Y. I. Kantor, and A. V. Sysoev. 1993. Foregut anatomy, feeding mechanisms, relationships and classifi- cation of the Conoidea (=Toxoglossa) (Gastropoda). Bul- letin of the Natural History Museum (Zoology) 59: 125— 170. Woodring, W. P. 1928. Miocene Mollusks from Bowden, Ja- maica. Part Il, Gastropoda and discussion of results. Car- negie Institute, Washington, Publication 85, 564 pp. MBL WHO! si INSTRUCTIONS TO AUTHORS I i THE NAUTILUS publishes articles on all aspects of the biology, a eee Ae and systematics of mollusks. Manuscripts describing original, unpublished research and review articles will ibe Bea rab er, Brief articles, not exceeding 1000 words, will be published as notes and do not require an abstract. Notices of interest to the mala- cological community will appear in a notices section. 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Box 1580 Sanibel, FL 33957 Y CONTENTS VIC t Volume 121, Number 4 December 21, 2007 ISSN 0028-1544 Thomas J. DeVries John Slapcinsky Robert Lasley Lennie Rotvit Jorgen Litzen Ase Jespersen Thomas Fox Eliane P. Arruda Osmar Domaneschi Jonata de A. Francisco José Carlos N. de Barros Donn L. Tippett Late Cenozoic Tegulinae (Gastropoda: Trochidae) from southern Peru... . Three new species of Paryphantopsis (Gastropoda: Pulmonata: Charopidae) from the Nakanai Mountains, New Britain, Papua: New Guineas... 5-06 eee eh ee dae Gees cee Seeds ase tbe Mysella gregaria new species, a bivalve (Galeommatoidea: Montacutidae ) to) D commensal with an intertidal burrowing sea anemone from North ‘Carolina, USAs «cis c0ch e008 eo gawas teehee ose h ew DEE HM Corbula tarasconii, a new species of Corbulidae (Bivalvia) from offshore: Brazil. p< ct a a eee odd Ghat oe wake eee baw ee ded Be hdd Two new gastropod species (Neogastropoda: Drilliidae, Turridae) from the western Atlantic Ocean . 0.0 163 182 190 STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION 1. Publication Title, THE NAUTILUS. 2. Publication No., 0028-1344. 3. Filing Date, October 1, 2007. 4. Issue Frequency, Quarterly. 5. No. of Issues Published Annually, Four. 6. Annual Subscription Price, US $80.00. 7. 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Total Number of Copies 430 430 B. Paid Circulation 1. Paid/Requested Outside-County Mail Subscriptions 298 308 2. Paid In-County Subscriptions 0 0 3. Sales Through Dealers and Carriers, Street Vendors, Counter Sales, and Other Non-USPS Paid Distribution 0 0 4. Other Classes Mailed Through the USPS 18 15 C. Total Paid and/or Requested Circulation 316 323 D. Free Distribution by Mail 1. Outside-County 12 12 2. In-County 0 0 3. Other Classes Mailed Through the USPS 0 0 E. Free Distribution Outside the Mail 0 0 F. Total Free Distribution 12 12 G. Total Distribution 328 335 H. Copies not Distributed 102 95 I. Total 430 430 J. Percent Paid and/or Requested Circulation 96% 96% THE NAUTILUS 121(4):163-1S1, 2007 Page 163 Late Cenozoic Tegulinae (Gastropoda: Trochidae) from outhern Peru Thomas J. DeVries! Burke Museum of Natural History and Culture University of Washington Seattle. WA 98195 US. A ABSTRACT Four new fossil tegulines (Gastropoda: Trochidae) are de- scribed from southern Peru [Chlorostoma quipua new species (late Miocene to late Pliocene), Intistoma pirqua new genus, new species (early Pliocene), Cantallocostoma panistostum new genus, new species (late Miocene to early Pliocene), Tegula (s.l.) masiasi new species (early to middle Miocene)], as are Pliocene and Pleistocene occurrences of the extant Chloro- stoma atrum (Lesson, 1830), C. luctuosum (d’Orbigny, 1841), Cantallocostoma quadricostatum (Wood, 1828), Agathistoma patagonicum (dOrbigny, 1835), T. (s.1.) melaleucos (Jonas, 1844), and T. (s.l.) tridentata (Potiez and Michaud, 1838). These data show that the Peruvian chlorostomine group is too ancient to be a Pliocene sister group to Caribbean-Atlantic agathistomines: indicate additional eastern Pacific groups of tegulines exist with roots reaching into the Miocene; and fur- ther demonstrate the success of A. patagonicum as a wide- spread and long-lived teguline in austral waters. A small radia- tion of Peruvian chlorostomines during the late Pliocene coin- cided with a molluscan mass extinction event in the Peruvian Faunal Province. Additional Keywords: Mollusk, Tegula, Chlorostoma, Agathi- stoma, Miocene, Pliocene, Cenozoic, Pisco Basin, Taxonomy INTRODUCTION Seven extant teguline species (Gastropoda: Trochidae) inhabit the cool coastal waters of the Peruvian Faunal Province. They are, according to their traditional nomen- clature, Tegula atra (Lesson, 1830), T. luctuosa (dOrbigny, 1841), T. ewryomphala (Jonas, 1844), T. ig- nota ae 1976, T. tridentata (Potiez and Michaud, 1838), T. quadricostata (Wood, 1528), and T. patagonica (dOrbigny, 1535) (Alamo and Valdivieso. 1997: Véliz and Vasquez, 2000). An eighth species, Tegula melalewcos (Jonas, 1544), a species from warmer waters of northern Peru, does appear rarely at southern Peruvian latitudes. This teguline fauna is as diverse as that of the warm-water Panamic Faunal Prov- * Mailing address: P.O. Box 13061, Burton, WA 98013 USA ince (Keen, 1971; Alamo and Valdivieso, 1997), although with mostly different species, and is more speciose than the teguline f auna of the Magellanic Faunal Province (four species), which is a subset of the Peruvian fauna (Forcelli, 2000). The geological record of tegulines from the Peruvian Faunal Province is meager. Tegula luctuosa, T. atra, and T. tridentata are listed in Herm’s (1969) study of Pliocene and Pleistocene mollusks from Chile. Tegula luctuosa and T. melaleucos were reported from upper Pliocene and Pleistocene beds of northern Peru (DeVries, 1986). Four new species of tegulines, consid- ered to have early to middle Miocene age (DeVries and Frassinetti, 2003: Finger et al., 2007), have been de- scribed from the Navidad Formation of central Chile by Nielsen et al. (2004). This paper documents four new fossil species and sev- eral previously known Recent species of tegulines in Neogene deposits of southem Peru, including the oldest leewn teguline from Peru, the early Miocene Tegula (s.l.) masiasi new species; creates two new genera of tegulines, Cantallocostoma, new genus, and Intistoma, new genus, each with a newly described Neogene fossil species in the Peruvian Faunal Province, Cantallocos- toma panistosum new species and Intistoma pirqua new species, and each having a modern representative in the eastern Pacific Ocean, the Peruvian C. quadricostatum and Californian I. aureotinctum (Forbes, 1852); adopts a full generic status for Tegula (Chlorostoma); and pro- vides evidence for a late Miocene origin of a western South American group of Chlorostoma species. GEOLOGY The late Cenozoic marine stratigraphy of southern Pe- ruvian forearc basins has been described by Dunbar et al (1990) and DeVries (1998). Teguline-bearing deposits crop out west of Nazca and throughout the Sacaco Basin (Fig. 1). These bioclastic conglomerates and sandstones which were assigned to the La Planchada and Pisco for- Page 164 THE NAUTILUS, Vol. 121, No. 4 Camana Basin’ 1 | 1 l 1 | Figure 1. Location of onshore portions of Cenozoic forearc basins in southern Peru. New fossil species of Tegula are from Cenozoic deposits west and south of Nazca. mations by Beaudet et al., 1976, and Muizon and DeVries, 1985, respectively, represent high-energy fore- shore and intertidal environments lying eleee to moun- tainous paleo-shorelines and quieter forediore and inner shelf environments lying hundreds to thousands of meters from paleo- shorelines defined by the beginning of the Andean foothills. MATERIALS AND METHODS Most Peruvian specimens described in this study were found by the author. Comparative material was studied at the Natural History Museuin of Los Angeles County, Los Angeles, California, USA (LACM). ‘Locality and sample descriptions are listed in the appendix. Lengths (L) and widths (W) are measured in millimeters, with dimensions of broken specimens enclosed by parenthe- ses. A non-standard orientation for photographs of some specimens has been necessary to reveal important char- acters. Some figured specimens were coated with ammo- nium chloride. ‘Type s and numbered specimens, includ- ing those figured, are deposited at the Orton Geological Museum, Ohio State University, Columbus, Ohio USA (OSU): the Departamento de Paleontologia de Vertebra- dos, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru (MUSM INY); and the Burke Museum of Natural History and Culture, Uni- versity of Washington, Seattle, Washington (UWBM). The prefixes “DV” refers to DeVries localities, “|M” to localities of J. Macharé (Instituto Geolé6gico Minero y Metalirgico, Lima, Peru), and “WIZ to localities of W. J. Zinsmeister (Purdue University, Indiana, USA). SYSTEMATICS Superfamily Trochoidea Rafinesque, 1815 Family ‘T Crochidae Rafine ssque, LS15 Subfamily Tegulinae Kuroda, Habe and Oyama, 1971 Genus Tegula Lesson, 1835 Type Species: Trochus elegans Lesson, 1835 (by monotypy) (= Trochus pellisserpentis Wood, 1828). Re- cent, Pacific coast of Central America. Remarks: Resolving the difficult subfamilial place- ment of Tegulinae wathib Trochoidea (Hickman and McLean, 1990: Bouchet and Rocroi, ‘ 2005) is beyond the scope of this paper. Tegula itself has been assigned to three different subfamilies over the past century, but most taxonomists now refer it to the undiagnosed Teg- ulinae Kuroda, Habe, and Oyama, 1971 (Hiclanen and McLean, 1990). Only one extant teguline species, Tegula pellisserpentis (Wood, 1828), pertains to Tegula a 'S5) (Keen, 1971). Its combination of foavares: (densely packed and heavily beaded spiral cords, tightly twisted columella, and numerous teeth extending from the col- umella across the apertural floor) is not seen in other species assigned to Tegula, so T. pellisserpentis is herein considered to be the monotypic representative of Tegula (s.s.), an opinion shared by J]. H. McLean (pers. comm., 2006). Taxa often considered as subgenera of Tegula, e.g., Chlorostoma and Agathistoma, are elevated to ge- neric status in this paper in accord with the practice of some authors (e.g., Ammold, 1907; Higo et al., 1999) and the opinion of J. HH. McLean (pers. comm., 2006). Genus Chlorostoma Swainson, 1840 Type Species: §Trochus argyrostomus Gmelin, 1791 (by subsequent designation, Herrmannsen, 1546). Re- cent, Japan. Remarks: Swainson (1840) erected Chlorostoma as a subgenus of Trochus Linnaeus, 1755, to include species with a “remarkably oblique” aperture, a very deep um- bilicus, oa one or two teeth on the outer (= lower part of inner ?) lip. He assigned two species s to Chlorostoma: Ti me (Chlorostoma) argyrostoma (= Tegula argyros- toma of authors) and Trochus (Chlorostoma) umbilicaris |= Gibbula umbilicaris (Linnaeus, 1758) |. Herrmannsen (1546) implicitly limited the definition of Chlorostoma by choosing T. argyrostoma as the type species. Examina- tion of LACM material from the western North Pacific Ocean [C. argyrostoma, C. lischkei (Tapparone-Canefri, 1874), C. rusticum (Gmelin, 1791), C. turbinatum (A. Adams, 1853), C. xanthostigma (A. Adams, 1853)] shows that adult chlorostomine umbilici can be either open or closed. In the view of Grant and Gale (1931), Chloros- toma should be further restricted to toothed species hav- ing ventricose whorls, a nacreous interior, and a dark purplish exterior. This diagnosis makes possible the iden- tification of Recent chlorostomine taxa from both sides of the North Pacific Ocean and in the eastern South Pacific Ocean. Chlorostoma pia (Lesson, 1830) (Figures 2-4, 7-9, 13) Trochus ater Lesson, 1830, vol. 2, pt. 1, p. 344, Mollusques, pl. 16, fig. 2: d Orbigny, 1S40: 409; Philippi, 1544, Abbildun- 5 T. DeVries, 2007 Page 165 Figures 2-4, 7-9, 13. Chlorostoma atrum (Lesson, 1830). 2. UWBM 97863, DV 1372-1, Recent, oblique spire view, width = 9.8 mm. 3. UWBM 97855, DV 395-1, Recent, basal view, width = 30.7 mm. 4. UWBM 97863, oblique basal view. 7. MUSM INV 126, DV 1252-1, early Pleistocene, oblique spire view, width = 27.1 mm. 8. UWBM 97855, i aalote view. 9. UWBM 97860, Paracas Hotel, Recent, lateral view, length = 22.2 mm. 13. UWBM 97860, oblique basal view, width = 23.4 mm. Figures 5,6. © Chlorostoma funebralis (A. Adams, 1855). UWBM 97562, south of La Jolla, California, Recent, width = 13.4 mm. 5. Oblique spire view. 6. Oblique basal view. Figures 10-12, 14-19. Chlorostoma luctuosum (d’Orbigny, 1841). 10. UWBM 97864, Paracas Hotel, Recent, lateral view, length = 20.2 mm. Il. UWBM 97865, Paracas Hotel, Recent, apertural view, length = 20.5 mm. 12. UWBM 97866, Hueco La Zorra, Peru, Recent, apertural view, length = 21.6 mm. 14. UWBM 97864, oblique basal view, width = 23.7 mm. 15. UWBM 97865, basal view, width = 24.4 mm. Figure 16. UWBM 975866, oblique spire view, width = 28.5 mm. 17. UWBM 97567, Lomas. Peru, Recent, oblique spire view, width = 26.4 mm. 18. UWBM 97867, oblique basal view. 19. UWBM 97868, Chile, Pleistocene, oblique basal view. width = 37.3 mm. Figures 20, 21. Chlorostoma ao aay (Jonas, 1844). UWBM 97871, DV 1599-1, Recent, length = 26.0 mm. 20. Apertural view. 21. Lateral view. Figures 22, 23. Chlorostoma ignotum (Ramirez-Bohme, 1976). UWBM 97872, Pellehue, Chile, Recent, width = 17.5 mm. 22. Oblique spire view. 23. Basal view. oo 24-26. Chlorostoma quipua new species. 24. UWBM 97873, DV 1254-Bal 6, late early Pliocene, basal view, width = 17.3 mm. Note faint protractive stripes on base. 25. MUSM INV 136, DV 571-1, syntype, late Miocene, oblique spire view, ak “170 mm. 26. UWBM 97873 oblique spire view. Page 166 gen und beschreibungen neuer oder wenig gekannter Conchylien, v. 1, p. 188, pl. 5, fig. 6; Philippi, 1846, Die Kreiselschnecken oder Trochoideen, p. 195, pl. 30, fig. 1; Hupé, 1854, p. 142, Malacologia, pl. 4, fig. 2. Monodonta atra Lesson.—Potiez and Michaud, 1838: 319, pl. 29, figs. 14, 15. Tegula atra Lesson.—Dall, 1909: 239, pl. 24, fig. 4; Carcelles and Williamson, 1951: 262; Aldea and Valdovinos, 2005: fig. SB. Morch, 1850: 20, Tegula (Chlorostoma) atra (Lesson, 1830).—Marincovich, 1973: 24, fig. 42; Alamo and Valdivieso, 1997: 13, fig. 25; Guzman et al., 1998: 35, fig. 22; Forcelli, 2000: 61, fig. 87; Véliz and Vasquez, 2000: 759, fig. 1B. Trochus moestus Jonas, 1844: 113; Philippi, 1846, Abbildungen und peschreibungen neuer oder wenig gekannter Con- chylien, v. 2, pl. 6, fig. 5; Philippi, 1 1846, Die Kre- isclsthnedlen’ oder Trochodecn. p. 199, Pr 30, fig. 2; Hupé, 1854: 147, Malacologia, pl. 4, figs. 3, 3a, 3b. Tegula moesta (Hupé, 1854).—D. all, 1909: 239: Alamo and Valdivieso, 1997: 14. Chlorostoma minor Morch, 1850: 20. Diagnosis: Shell width about 40 mm; last whorl broadly rounded, including shoulder; keels lacking. Um- bilicus of juvenile and adult shells white, closed; meni cus with two well exposed spiral cords, the adaxial cord terminating in a columellar tooth. Material Examined: UWBM 97855, DV 398-1, Re- cent, L = 25.0, W = 30.7; UWBM 97856, DV 398-1, lot of 2; UWBM 97857, DV 1252-1, L = 11.8, W = 18.6; UWBM 97858, DV 1418-1, latest Pliocene, L = (22.2), W = 30.2; UWBM 97859, DV 463-1, late Pleistocene, lot of 2; UWBM 97860, Paracas Hotel, Recent, L = 22.2; W = 23.4; UWBM 97861, Ipun, Chile, Recent, lot of 2; UW BM 97863, DV 1372-1, Recent, L = 8.0, W = 9.8; MUSM INV 126, DV 1252-1, early Pleistocene, L = (19.2), W = 27.1; MUSM INV 127, DV 1418-1, L = 31.4, W = 36.6; MUSM INV 128, DV 463-1, lot of 2. Occurrence: Late Pliocene to middle Pleistocene: southern Peru to southern Chile. Late Pleistocene: southern Peru, Chile, southern Argentina. Recent: northern Peru to southern Chile, southerm Argentina (G. Pastorino, pers. comim., 2002). Remarks: Specimens of Chlorostoma atrum can ex- ceed 40 mm in width and are generally smooth-shelled and purple-black, either entirely or dorsally, only. The last whorl is always broadly rounded; it lacks the keeled spiral cords present on specimens of C. luctuosum. The base on some specimens of C. atrwm has weak spiral threads; the spire of some also has one or more narrow spiral grooves that produce as many as 15 intervening low broad spiral cords (Figure 7), not unlike the spiral sculp- ture of the Californian C. funebralis (A. Adams, 1855) (Figures 5, 6), The adult shell of Chlorostoma atrum is usually dis- tinguished from that of other Peruvian chlorostomines by its closed umbilicus. [G. Collado (pers. comm., 2005) notes that juveniles of C. luctwosum and adults of the small Tegula (s.1.) tridentata occasionally have closed umbilici.| The white umbilical area on specimens of C. atrum has two spiral cords. A white inner cord rises from THE NAUTILUS, Vol. 121, No. 4 beneath the umbilical callus and terminates on the edge of the columella as a thickened tooth. A weakly devel- oped white outer cord traces the boundary of the um- bilical area and becomes flattened on the columella, not quite protruding far enough to produce a tooth. Rarely, one or two weak spiral spurs develop between the two umbilical cords in a nacreous area that lies adaperturally of a thin, glossy, umbilical veneer. They, too, do not extend far enough to produce columellar teeth. Chlorostoma luctuosum (VOrbigny, 1841) (Figures 10-12, 14-19) Trochus luctuosus d’Orbigny, 1841, v. 5, p. 409, pl. 76, figs. 16-19; Philippi, 1846, Die Kreiselschnecken oder Tro- choideen, p. 153, pl. 25, figs. 4, 5; Hupé, 1854: 143. Tegula luctuosa Orbigny.—D Dall, 1909: 239: DeVries, 1986: 512, pl. 27, figs. 3, 4; Guzman et al., 1995: 36, fig. 23; Véliz and Vasquez, 2000: 762, fig. 1D: Alamo and Valdivieso, 2000: 14; Aldea and Valdovines, 2005: fig. SE. Diagnosis: Shell width to 35 mm. Last whorl with one to three spiral cords or keels. Adult umbilicus open; ju- venile umbilicus usually open; umbilicus with two well exposed umbilical spiral cords, the adaxial cord terminat- ing in a columellar tooth. Material Examined: OSU 37596, DV 240-23, latest Pliocene, L = 19.5, W = 23.9: UWBM 97864, Paracas Hotel, Recent, L = 20.3, W = 23.7; UWBM 97865, Pa- racas Hotel, Recent, L = 20.5, W = 24.4: UWBM 97866, Hueco La Zorra, Recent, L = 21.6, W = 28.5; UWBM 97867, Lomas dump, Recent, L = (20.6), W = 26.4: UWBM 97868, W]Z 345, Chile, Pleistocene, L = 26.8, W = 37.3; UWBM 97869, DV 382-1, Pleistocene, L = (20), WwW. 7 98.4: UWBM 97870, WIZ 345, L = (35.0), W = 43.1; MUSM INV 129, DV 382-1, L = 20.6, W = 31.1; MUSM INV 130, JM 8220, Pleistocene, lot of 2. Occurrence: Late Pliocene to upper Pleistocene: northern Peru to Tongoy, central Chile. Recent: Galapa- gos Islands to Concepcion, central Chile (southern limit from LACM collections). Remarks: Specimens of Chlorostoma luctuosum are large and purple-black, either entirely or dorsally, only. Adult specimens are generally distinguished from speci- mens of C. atrum by having an open cambaeus and from both C. atrum and C. euryomphalum by having one to three primary spiral cords or keels: one near a base of the whorl, forming the periphery (Figure 12); another about one quarter of the distance anteriorly from suture to suture (Figure 12); and a third occasional] y dev eloped just anterior to the periphery (Figure 19). Some speci- mens of C. luctuoswm are also cov ened with tertiary spiral threads (Figures 17, 1S). The thin umbilical veneer and columellar teeth are identical to those on specimens of C. atrum, as are the umbilical cords, except that they are exposed coiling deep into the umbilicus. Chlorostoma euryomphalum (Jonas, 1844) (Figures 20, 21) Trochus euryomphalus Jonas, 1844; 113; Philippi, 1844, Abbil- dungen und beschreibungen neuer oder wenig gekannter T. DeVries, 2007 Page 167 774250 W x sxOIH NVOIANVNVd SS Aguada de Lomas Pacific Ocean e@ DV _ locality—sample 0 Skm a SCALE Contour interval is 100 m Figure 41. new species. Type locality (DV 571-1) of Chlorostoma quipua Conchylien, v. 2, p. 27, pl. 6, fig. 4; Philippi, 1846, Die Kreiselschnecken . Wi schotdeen, p 15h aL 25, fig. 7. Tegula euryomphala (Jonas, 1844).—Carcelles and W illiamson, 1951: 262. Tegula euryomphalus [sic] (Jonas).—Dall, 1909: 239; Alamo and Valdivieso, 1997: 14. Tegula euryomphala (Jones, 1844) [sic] —Guzman et al., 1998: 36, fig. 25; Véliz and Vasquez, 2000: 762, fig. LE; Aldea and Valdovinos, 2005: fig. SC. Trochus kieneri Hupé, 1854, p. 144, Malacologia, pl. 4, figs. 1, la. 1b. Diagnosis: Shell width to 35 mm. Last whorl broadly rounded. Umbilicus white, open; umbilicus with two we il] exposed spiral cords, the adaxial cord terminating in a columellar tooth. Material Examined: U\WBM 97871, DV 1599-1, Re- cent, L = 26.0, W = 29.9. Occurrence: Late Pleistocene: Northern to central Chile. Recent: Southern Peru to central Chile. Remarks: Specimens of Chlorostoma euryomphalum are large, purple-black, and characterized by a broad open umbilicus and broadly rounded whorls. They differ from specimens of C. atrum. which have a closed umbi- licus, and C. luctuosum, which have one or more angular spiral cords or keels. On some specimens of C. luctio- sum, however, including Recent Peruvian and Chilean examples from LACM collections, Pleistocene Chilean specimens from WJZ collections, and upper Pliocene Pe- ruvian specimens from northern Peru, the spiral cords are so weak that assigning the material to C. luctuosum or C. euryomphalum is problematic. Chlorostoma ignotum (Ramirez-B6hme, 1976) (Figures 22, 23) Tegula ignota Ramirez-Bohme, 1976: 3, figs. 1-6; Forcelli, 2000: 61, fig. 8S; Véliz and Vasquez, 2000: 762, fig. 1F; Aldea and V idoanos 2005: fig. SD; Collado ond Brown, 2005: 131. Diagnosis: Shell width to 30 mm. Outer layer slate colored. Sculpture consists of several well developed, un- beaded, primary spiral cords. Umbilicus open. UWBM 97872, La Rinconada, 11.1, W = 17.5; UWBM 29.4, W = 32.6. Material Examined: Pelluhue, Chile, Recent, L = 97906, Ipun, Chile, Recent, L = Occurrence: Recent: central to southern Chile. Remarks: = Chlorostoma ignotum is known only as a Recent species from Chile (e.g., Collado and Brown, 2005). Specimens of C. ignotum differ from those of other extant chlorostomines of the Peruvian Faunal Province in two significant respects: they lack the purple- black outer shell jan er of C. atrum, C. luctuosum, and C. euryomphalum, being rather slate colored, and they have numerous, prominent, well-differentiated primary and secondary spiral cords between the periphery and suture and to a lesser extent on the base. The well-exposed umbilical cords, thin umbilical veneer, and columellar teeth are identical, however, with those of other species of South aac Chlorostoma, and the strong spiral cords (Figure 22) are like those seen on rare specimens of early Plioc ‘ene C, quipua new species (Figures 25, 26; see below) Chlorostoma quipua new species (Figures 24—40) Diagnosis: Adult whorls purple-black dorsally; spire and base usually light brown, commonly with dark brown protractive stripe s or mottling. Umbilicus narrow, open. Description: Shell up to 30 mm wide. Spire angle about S0 degrees. Periphery near base, sharply rounded to slightly angul uw. Sutures appressed. Protoconch un- known: te Jeoconch with five flat-sided to slightly convex whorls. Axial sculpture absent or rarely with rugose pro- tractive ribs. Thin growth lines strongly prosocline. Spiral sculpture of 20 muted spiral thre ads: poste rior to periph- ery; rarely with three to five spiral grooves separating four to six broad low spiral cords or mthioul spiral sculp- ture. Twenty to 30 evenly spaced spiral threads on base of juvenile specimens, muted or obsolete on adult whorls. Outer shell layer purple-black on adult whorls, tan or light brown on spire whorls and base. Protractive and rarely retrotractive wrinkled brown stripes usually Page 168 THE NAUTILUS, Vol. 121, No. 4 Figures 27-40. Chilorostoma quipua new species. 27. MUSM INV 131, DV 571-1, syntype, spire view, width = 25.4 mm. 28. MUSM INV 133, DV 1635-2, early Pliocene, basal view, width = 16.0 mm. 29. UWBM 97880, DV 1635-2, basal view, width = 16.6 mim. 30. UWBM 97879, DV 1635-2, basal view, width = 18.5 mm. 31. UWBM 97590, DV 1598-1, early Pliocene, oblique spire, width = 11.2 mm. 32. UWBM 97883, DV 1254-1, Pliocene, oblique basal view, width = §.9 mm. 33. UWBM 97876, DV 571-1, syntype, apertural view, length = 22.7 mm. 34. UW BM 97880, oblique spire view. 35. MUSM INV 133, apertural view. 36. UWBM 97581, DV 1029-1, early Pliocene, spire view, width = 17.4 mm. 37. UWBM 97879, oblique spire view. 38. MUSM INV 135, DV 1029-1, oblique spire view, width = 11.3 mm. 39. UWBM 97878, DV 809-1, Pliocene, spire view, width = 5 809-1, basal view, width = 19.0 mm. 2.1 mm. 40. MUSM INV 132,DV present on base and less often on spire. Inside edge of Type Material: (All DV 571-1, all syntypes) UWBM outer lip smooth. Umbilicus open, narrow; umbilicé al ve- 97876, L = 22.7, W = 27.3; UWBM 97877, L = (17), neer thin. Columella with thick inner tooth at end of W = (25.1); MUSM INV 131, L = (18.9), W = 28.4; well-exposed white umbilical cord. Weak outer MUSM INV 136, L = (10.2), W = 17.0. tooth adjacent to floor of aperture at end of thin spiral Other Material Examined: UWBM 97873. DV1254- cord following outer margin of umbilical area. Small pa- Bal 6, late early Pliocene: 1, = (11.0): W = 17.3: UWBM rietal flange barely ove thanging umbilicus. 97874. DV 1254-Bal 10, late Pliocene, L = (4.8), W = Type Locality: DV 571-1, Alto Grande, about one km 10.4: UWBM 97875, DV 1254-Bal 10, L = (14.1), W = south of intersection with abandoned paved road to San 24.4: UWBM 97878, DV SO9-1, Pliocene, L = 15.4, W = Juan de Marcona, on south-facing hillside west of Pan- 22.) pe 97879, DV 1635-2, early Pliocene, L american Highway; one of several shell banks of the 12.2. W = 18.5: UWBM 97880, DV 1635-2, L = (11.3), Pisco Formation (Figure 41). Locality inaccurately re- W = 16.6; UWBM 97881, DV 1029-1, early Pliocene, ferred to as El Jahui w in Muizon and DeVries (1985). L = (12.3), W = (17.4); UWBM ae DV 1029-1. L = 15°26'57"S, 74°52'06"W (Acari 1:100,000 quadrangle), (12), W = 16.7; UWBM 97883, DV 1284-1, Pliocene, Middle upper Miocene, L = (5.2), W = 8.9; UWBM 97890, DV 1598-1, early T. DeVries, 2007 Page 169 Pliocene, L = 8.4, W = 11.2; MUSM INV 132, DV SO9-1, = gr W = (19.0); MUSM INV 133, DV 1635-2, L = 11.2, W = 16.0; MUSM INV 134, 1635-2 lot of 2; MUS INV 135, DV 1029-1, L = 6.9, W = 11.3 Occurrence: Middle late Miocene to early late Pliocene: southern Peru. Etymology: “Quipua,” Latinized version of “quipu,” Inca counting device of braided and knotted strings, evoked by the wrinkled brown stripes on the base and spire of this species. Remarks: Specimens of Chlorostoma quipua difter from those of C. euryomphalum and C. luctuosum by having a narrower cabied area, smaller parietal f flange, and protractiv' e brown stripe Ss. Specimens of C. quipua lack the keeled spiral cords of C. luctuosim and closed umbilicus of C. atrum. Some specimens of C. quipua, both Miocene and Pliocene, have broad spiral cords (Fig- ure 25) like those seen on the juvenile whorls of some specimens of C. atrum (Figure 7). A single specimen from upper Pliocene beds above Playa Huacllaco ( Figure 26) has spiral cords as pronounced as the raised spiral cords on specimens of the modern Chilean C. ignotuim (Figure 22). Some lower Pliocene specimens near Yauca (Figure 31) have coarse protractive axial ribs like those on some Asian chlorostomines. Specimens of Chlorostoma quipua superficially re- semble those of C. gallina (Forbes, 1852), a Pliocene-to- Recent species from California and Baja California (Grant and Gale, 1931; McLean, 1978), and C. rugosiwm (A. Adams, 1853), a Recent species from the Gulf of California (Keen, 1971). Specimens of all three species have some degree of purple-black color and protractive stripes on the spire and/or base. Specimens of C. gallina and C. rugosum, however, are more ventricose laterally and bz sally and have weak to prominent protractive axial ribs and stripes posterior to the base. Specimens of C, gallina usually have a closed umbilicus. Shells of Chlorostoma quipua are found in upper Mi- ocene beach deposits near Alto Grande (DV 571-1; see Muizon and DeVries. 1985) with specimens of Chorus frassinetti DeVries, 1997, and Acanthina obesa DeVries, 2003 (DeVries, 1997, 2003). Lower Pliocene specimens of C. quipua occur together with specimens of the mu- ricid gastropods, Concholepas kieneri Hupé, 1854; Xan- thochorus ochuroma DeVries, 2005; and Herminespina saskiae DeVries and Vermeij, 1997: and the turbinid gas- tropod, Prisogaster mcleani DeVries, 2006 (DeVries, 2005, 2006; DeVries and Vermeij, 1997) Genus Cantallocostoma new genus Type species: Trochus quadricostatus \Vood, 1828. Recent, Peru and Chile. Diagnosis: White to brown outer shell layer. Three to five beaded primary spiral cords. U mbilicus open, broad, Two adaxially situated spiral umbilical cords terminating in columellar teeth. Parietal wall vertical, without pari- etal flange overhanging umbilicus. Description: Shell wp to 35 mm in diameter. Whorls ventricose to quadrate; periphery weakly bicarinate. Spi- ral sculpture of three to five primary spiral cords bro- ken into beads; interspaces with two to five continuous or weakly beaded tertiary threads. Base with four to five beaded or non-beaded primary spiral cords and in- tervening secondary cords and tertiary threads. Um- bilical area white, tabulate, sh: ply de fined, with margin of umbilical area flaring towards aperture as steep- ly inclined wall. U salslious open, broad, with two spiral umbilical cords situated adaxiz ily, the innermost thicker; each cord terminating in columellar tooth. Third tooth sometimes present at base of columella. Um- bilical veneer variably developed. Columella thin, upright, without parietal flange overhanging umbili- cus. Floor of aperture with le -dge but without teeth. In- ner lip sometimes with four to six closely spaced low teeth. Occurrence: Late Miocene to Pleistocene: southern Peru. Recent: northern Peru to Chile. Etymology: “Cantalloc,” site near Nazca, Peru, where subterranean aqueducts are reached from ground level by pre-Incaic stonewall-lined spiral paths that resemble the spiral umbilical cords of this genus. Remarks: Specimens of Cantallocostoma differ from those of Asian, Californian, and Peruvian Chlorostoma by having beaded spiral cords and two adaxially situated amabileeal spiral cords. They differ from specimens of Intistoma new genus, by having closely spaced beads and lacking a subsutural band of well-deve loped protractive nodes, Specimens of Cantallocostoma fer from teg- ulines traditionally assigned to Agathistoma by lacking hallmarks of that genus: “narrow open umbilicus, a smooth or finely beads d spiral sculpture, and a varie- gated surface coloration” (Olsson and Harbison, 1953: 351). Cantallocostoma quadricostatum (Wood, 1825) (Figures 42-45, 47-49) Trochus quadricostata Wood, 1828: 16, pl. 5, fig. 16. Trochus quadricostatus Wood.—Philippi, 1546, Die krei- selschnecken oder Trochoideen p. 154, pl. 25, fig. 6. Tegula quadricostata (Wood, 1828).—Véliz and Vasquez, 2000 759, fig. LA; Aldea and Valdovinos, 2005: SF. Tegula quadricostata Gray.—Dall, 1909: 240; Alamo and Valdivieso, 1997: 14. Monodonta catenifera Potiez and Michaud, 1838: 315, pl figs. 12-13. Trochus torulosus Philippi, 1543, bungen neuer oder wenig gekannter Conchylien, v. 1, pl 2. fig. 12. Abbildungen und Beschrei- Diagnosis: Low spire, white to cream-colored with purple along edges of sculptural elements. Sculpture of three to five prominent primary beaded vee cords. Material Examined: UWBM 97S8S4, DV 13 e cent, | (14.4), W = 23.9: UWBM 978585, D\ y= : ; L = (13.3), W = (20.8); UWBM 97886, DV 1713-1, early 2-1, R 372-] Page 170 THE NAUTILUS, Vol. 121, No. 4 Figures 42-45, 47-49. Cantallocostoma quadricostatum Wood, 1828. 42. UWBM 97884, DV 1372-1, Recent, oblique spire view, width = 23.9 mm. 43. UWBM 97886, DV 1713-1, early Pleistocene, apertual view, width = 21.5 mm. 44. MUSM INV 137, DV 1355-1, late Pliocene, basal view, width = 16.0 mm. 45. UWBM 97854, oblique basal view. 47. UWBM 97885, DV 1372-1, oblique lateral view, width = 20.8 mm. 48. UWBM 97887, DV 1355-1, oblique spire view, width = 18.1 mm. 49. UWBM 97886, oblique basal view. Figures 46, 50-54. Cantallocostoma panistostum new species. 46. MUSM INV 141, DV 1635-2, early Pliocene, oblique spire view, width = 14.3 mm. 50. UWBM 97891, DV 571-1, late Miocene, oblique spire view, width = 14.9 mm. 51. UWBM 97889, DV 1598-1, syntype, oblique lateral view, width = 33.7 mm. 52. UWBM 97889, oblique basal view. 53. UWBM 97889, apertural view. 54. MUSM INV 139, DV 1598-1, syntype, basal view, width = 24.9 mm. Figures 55-57. Intistoma pirqua new species. UWBM 97892, DV 470-1, syntype, early Pliocene, length = 38.9 mm. 55. Lateral view. 56. Apertural view (lighting from upper right). 57. Oblique basal view. Figures 58,59. — Intistoma aureotinctum (Forbes, 1852). South of La Jolla, California, Recent. 58. UWBM 97596, lateral view, length = 20.0 mm. 59. UWBM 97897, oblique basal view, width = 17.8 mm. Figures 60-63. Agathistoma patagonicum (VOrbigny, 1535). 60. UWBM 97893, Argentina, Recent, oblique spire view, width = 15.4 mm. 61. UWBM 97893, oblique basal view. 62. UWBM 97895, DV 1032-2, late Pliocene, lateral view, width = 12.7 mm. 63. UWBM 97895, oblique spire view, T. DeVries, 2007 Pleistocene, L = (14.5), W = 21.5: haley 97SS7, DV 1355-1, late Pliocene, L = — V = 18.1; UWBM OE? DV 1355-1], L, = (10.0), ae 1; MUSM INV , DV 1355-1, L = (8.9), W = ie MUSM INV 138, DV 1355-1, L = 14.1, W = (19). Occurrence: Late Pliocene: southern Peru. Recent: northern Peru to Chile. Remarks: The number of primary beaded spiral cords on the last whorl of Cantallocostoma quadricostatum var- ies between three and five. Most modem specimens have two widely spaced primary spiral cords on the anterior half of the whorl and two closely spaced primary spiral cords adjacent to the posterior suture (Figure 42). Some specimens have an additional primary spiral cord be- tween the two anterior spiral cords (Figure 47); other specimens have one of the two posteriormost primary spiral cords missing (Figure 43). The six known late Pliocene specimens from southern Peru (Figure 45) and single early Pleistocene specimen (Figure 43) have three primary spiral cords. Cantallocostoma panistostum new species (Figures 46, 50-54) Diagnosis: Exterior uniformly pale brown; spiral sculpture of three beaded spiral cords. Basal spiral cords bunched towards umbilical area. i Shell nearly 35 mm in diameter. Spire angle about 70 degrees. Periphery at base, sharply rounded. Sutures impressed. Protoconch unknown. Te- leoconch with five quadrate whorls. Axial sculpture ab- sent except for intermittently rugose prosocline growth lines. Spiral sculpture of * closely beaded spiral Sond at base and two posterior spiral rows of more widely spaced beads, about 25 per whorl, bracketing shoulder. Poste- rior beads sometimes slightly protractively elongate. In- terspaces rarely with beaded secondary i cords: usu- ally with wavering tertiary spiral threads. Base weakly convex, with six to ten closely spaced spiral cords adja- cent to umbilical area. Outer shell layer pale brown, without color pattern. Inner edge of outer lip smooth. Umbilicus open, wide, with two adaxially situated spiral cords. Umbilical veneer thin. Innermost umbilical cord prominent, second cord thin; each terminates in col- umellar tooth. Third tooth at base of columella nearly obsolete. Columella thin; parietal flange absent. Type Locality: DV 1595-1. eres along Panameri- can Highway, shell beds exposed along descent from north into Rio Yauca valley (Figure 64). 15°39'49"S, 74°31'50"W (Yauca 1:100,000 quadrangle). Lower Pliocene. Type Material: (DV 1595-1, all syntypes) UWBM 97889, DV 1598-1, L = (25.2), W = 33.7: UWBM 97890. © DV Locality-samples (e) Town or farm 0) 5km SCALE Contours in meters Figure 64. Type locality (DV 1598-1) of Cantallocostoma panistostum new species and other teguline localities between Sacaco and Yauca. DV 1598-1, L = 8.4, W = 11.2; MUSM INV 139, DV 1598-1, L = (13.7), W = 24.9: MUSM INV 140, Pan- american roadcut at Yauca, early Pliocene, L = (17.9), V = 33.3. Other Material Examined: UWBM 97891, DV 571- 1, late Miocene, L = 10.4, W = 14.9; MUSM INV 141, DV 1635-2, early Pliocene, L = 8.8, W = 14.3. Occurrence: Pliocene: south- ern Peru. Late Miocene to early Etymology: “Panis,” Latin noun meaning “bread,” and “tostum,” Latin neuter past participle- adjective me aning “toasted,” referring to the bread-crust color of this spe- cles. Remarks: The light brown color of the outer shell layer on specimens ‘of Cantallocostoma panistostum re- sembles that of specimens of Tegula hemphilli Oldroyd, 1921, a late Pliocene-to-Pleistocene species from Cali- fornia (Grant and Gale, 1931). Specimens of T. hemprhilli and numerous other Miocene and Pliocene Californian teguline species with similar coloration lack the two um- bilical spiral cords close to the axis and are covered by numerous closely spaced primary spiral cords, none of which are beaded. Specimens of Cantallocostoma panistostum are found in upper Miocene nearshore sandstones with Chloros- toma quipua; Chorus frassinetti DeVries, 1997; Acan- thina obesa; and Xanthochorus stephanicus DeVries, 2005; and in lower Pliocene cobbly bioclastic gravels as- sociated with the mouth of the paleo-Rio Yauca with disarticulated valves of an undescribed Anadara species, venerid bivalves, Chlorostoma quipua, Xanthochorus ochuroma, and Concholepas nodosa Moricke, 1896. Genus Intistoma new genus Type species: Trochus aureotinctus Forbes, 1852 Pleistocene to Recent, California. j0e wd) Page 2, THE NAUTILUS, Vol. 121, No. 4 Diagnosis: Spiral sculpture of subsutural band of thick protractive nodes and peripheral and sub-peripheral pri- mary spiral cords, Base with three thick, primary spiral cords. Umbilicus open. Description: Shell up to 45 mm wide, spire angle about 75 degrees. Whorls four to five in number, quad- rate to carinate; sutures weakly impressed. Protoconch unknown. Sculpture of thick rounded protractive axial ribs intersecting with an equally thick spiral cords, pro- ducing a broad subsutural spiral band of elongate pro- tractive nodes, a near-basal peripheral band of stubby protractive nodes more numerous than nodes in the sub- sutural band, and a sub-peripheral primary spiral cord with little axial modification. Tertiary threads sometimes present; often corrugated by slightly raised strongly ob- lique lamellar growth lines. Umbilicus open. Columella thin; parietal flange erect to slightly overhanging umbi- licus: parietal callus small. Umbilical veneer thick, cov- ering all but wedge-shaped adapertural portion of um- bilical wall. Umbilical spiral cord submerged in umbilical wall, emergent terminally as prominent columellar tooth. Smaller second columellar tooth sometimes present abaxially adjacent to first tooth. Etymology: “Inti,” the Inca sun god, with a nod to California's sunshine and the sunset-orange color inside the umbilicus of the type species, Intistoma aureotinc- tum. Occurrence: Late Miocene or early Pliocene: south- ern Peru. Early Pleistocene to Recent: California. Remarks: The new genus, Intistoma, is proposed for two very similar species: the Pleistocene-to-Recent Cali- ornian Intistoma aureotinctum and the early Pliocene Peruvian I. pirqua new species. Specimens of both spe- cies differ from those of nearly all other teguline taxa by possessing three thick primary spiral cords on the b: ise, rather than cords that are more numerous and thinner. Specimens of Intistoma additionally differ from those oroperly assigned to Agathistoma by ‘lac ‘king teeth on the loor of the aperture aud the inner edge of the outer lip and by lacking closely spaced be aded primary spiral cords. Intistoma aureotinctum has been considered the ex- tant representative of a lineage of Californian Neogene tegulines (Addicott, 1970) that includes the early Mi- ocene Tegula dalli arnoldi Addicott, 1970, the late Mi- ocene Tegula nashae Clark, 1915, and Pliocene Tegula hemphilli Oldroyd, 1921. Specimens of fossil ‘aliforniz m spec ies do have a subsutural band of e longate protrac tive nodes, as do specimens of Intistoma, and some have thick peripheral spiral cords, but none have the distinc- tive intistomine combination of thick basal spiral cords and tertiary spiral threads across the entire surface of the W he rls Intistoma pirqua new species (Figures 55-57) Diagnosis: Shell large, weakly bicarinate, with poste- rior row of protractive nodes. Base of shell with three broad spiral cords. Umbilicus open. Description: Shell large, width more than 40 mm; spire angle about 60 degre ees. Whorls ventricose, weakly bicaraate: sutures deeply impressed. Protoconch un- known; early spire whorls missing; teleoconch of at least three whorls. Last whorl with sculpture of about 18 elon- gate protractive ribs on posterior half of whorl; with about 21 circular to slightly protractively elongate nodes at periphery, two- ‘hinds the distance from suture to base: and with a continuous low broad spiral cord at edge of base. Base with three low broad spiral cords, innermost cord bordering umbilical area and twice as wide as other two cords. Faint traces of secondary spiral cords in in- terspaces laterally and basally. Aperture oblique, outer lip and colabral growth lines’ moderately prosocline (40 degrees). Wrbilieus open. Columella with at least one tooth, directed basally; anterior portion par tly excavated, partly missing. Floor of aperture partly missing; no teeth evident. Type Locality: DV 470-1, above rocky road from Ha- cienda Tunca to Quebrada Huaricangana; hillside of brown sandstone (Figure 65). 14°56" S, 75°09! W (Palpa 1:100,000 quadrangle). Upper Miocene or lower Pliocene. Type Material: (DV 470-1, syntypes, late Miocene or © Towns or villages @ DV locality-samples 0 5 10 km — SCALE Contour interval is 500 m. PUERTO’ < 9 CABALLAS . £ " va MearicahO Ze \ on ‘Cerro Huar icangana —15°00'S Figures 65. new species. Type locality (DV 470-1) of Intistoma pirqua T. DeVries, 2007 >I ww Page | early Pliocene) nae Aig L = (388.9), W MUSM INV 142, L 2), W = (42). Occurrence: ern Peru. Late Miocene to early Pliocene: south- Etymology: “Pirqua,” Latinized version of “pirqa,” Quechua word for “wall,” referring to the similarity of this species’s sculpture and Incaic stone walls. Remarks: The type specimens of Intistoma pirqua closely resembles specimens of I. aureotinctum (Figures 58.59), diff ffering principally by being twice the size in all dimensions and by having better dev veloped nodes on the peripheral spiral cord. The specimens of I. pirqua were found together with specimens of Chlamys simpsoni (Philippi, 1887) and Panopea coquimbensis (dOrbigny, 1842), both species from lower Pliocene beds in Chile (Herm, 1969) and southern Peru (Muizon and DeVries, 1985). Genus Agathistoma Olsson and Harbison, 1953 Type Species: Trochus viridulus Gmelin, 1791 (by original designation). Recent, Caribbean and northeast- ern South America. Remarks: One of two Recent specimens of Agathis- toma patagonicum from Argentina (UWBM 97893) lacks an open umbilicus (Figure 61) , as do some specimens of the Pliocene Sierra Laziar outcrops in Argentina (Iher- ing, 1907), suggesting either that the character is not diagnostic for all species of Agathistoma (Olsson and Harbison, 1953) or that the species in question might not be a member of the Agathistoma group. Agathistoma patagonicum (d’Orbigny, 1835) (Figures 60-63) Trochus (Monodonta) patagonicus “i Orbigny, 1535, vol. 3(4), p. 155; d’Orbigny, 1840, vol. 5(3), p. 408, pl. 55, figs. 1-4. Neomphalius patagonicus ( (Orb.). ). —thering, 1907: 400. Tegula patagonica Orbigny.—Dall, 1909: 240; Alamo and Valdivieso, 1997: 14; Forcelli, 2000: 62, fig. 89. Tegula (Agathistoma) patagonica (dOrbigny, 1835).—Rios, 1985: 20, pl. 9, fig. 77; Del Rio, 1998: 27, pl. 1, figs. 16-17. Trochus corrugatus Philippi, 1844, Abbildungen und Beschire 2 bungen neuer oder wenig gekannter C onchylie n,v. 1, p. 67, pl. 2, fig. 7. Trochus fuscesens Philippi, 1844, Abbildungen und Beschrei- bungen neuer oder wenig gekannter Conchylien, v. 1, p. 92, pl. 3. fig. 8 Trochus ae gnyana Pilsbry, 1900: 110; Carcelles, 1945: 38, pl. 1, figs. 6, 7, 12-15. Material Examined: UWBM 97893, Cabo dos Bahias. Chubut Province, Argentina, Recent, L = 11.2, W = 15.4: UWBM 97894. Cabo dos Bahfas, Chubut Province. Argentina, Recent, L = 11.6, W = 14.6; UWBM 97895, DV 1032-2. late Pliocene, L = (9), W = 12.7. Diagnosis: closely spaced beaded primary spiral cords with interca- Shell under 20 mm wide. Sculpture of lated secondary cords; beading sometimes obsolete. Col- umella with three teeth: ee ae open or closed. Occurrence: Late early to middle Miocene: Argentina (Ihering, 1907). Late Pliocene: southern Peru. Recent: northern Peru to Chile, southern Brazil to Argentina. Remarks: A single incompletely preserved specimen of an agathistomine was found between Yauca and Chala in bioclastic deposits just below the highest marine ter- race at 200 meters above sea level. Associated taxa that are either locally or entirely extinct |Prisogaster valenciai DeVries, 2006; Acanthina triangularis DeVries, 2003; Chorus giganteus (Lesson, 1830); Concholepas camerata DeVries, 2000; Xanthochorus xuster DeVries, 2005] are indicative of a late Pliocene age (DeVries, 1997; 2000; 2003: 2005: 2006). The closely spaced beaded spiral cords on the south- ern Peruvian agathistomine resemble those on speci- mens of Agathistoma verrucosum McLean, 1970, and A. pictum McLean, 1970, Panamic species which pre ssently range as far south as northern Peru (Alamo and Valdivieso, 1997), but the base of the Peruvian Pliocene specimen is not as flattened as it is on specimens of the northern Peruvian species and the spiral cords are more closely spaced, suggesting an assignment to A. patagoni- cum. Genus or Subgenus indeterminate Tegula (s.1.) tridentata (Potiez and Michaud, 1838) (Figures 66, 65) Monodonta tridentata Potiez and Michaud, 1838, vol. 1, p. 321, pl. 29, figs. 16-17. Trochus tridentatus Potiez and Michaud.—Philippi, 1546, Die Kreiselschnecken oder Trochoideen, p. 153, pl. 25, fig. 3. Tegula tridentata (Potiez and Michaud).—Dall, 1909: 176; Car- celles and Williamson, 1951: 262; Herm, 1969: 91; Aldea and Valdovinos, 2005: fig, SG. Tegula (Chlorostoma) tridentata (Potiez and Michaud, 1838).—Marincovich, 1973: 24, fig. 43; Véliz and Vasquez, 2000; p. 761, fig. 1C; Guzman et al., 1998: 36, fig. 24; Forcelli, 2000: 90, fig. 60. Trochus tridens Hupé, 1854: 145, Material Examined: UWBM 97898, DV 1372-1, Re- cent, L = 7.0, W = 9.9: UWBM 97899, DV es Re- cent, lot of 5: UWBM 97900, DV 381-5, middle Pleis- tocene, L = (11.7), W = 14.0. Occurrence: Middle Pleistocene: southern Peru to central Chile. Recent: northern Peru to southern Chile. Remarks: Tegula tridentata is a small, high-spired, purple-black species with an excavated white base. The exterior is usually smooth, but some specimens have low broad primary spiral cords on juvenile whorls. A narrow open aanbilicis is bordered by a white columella with three teeth. The two teeth closest to the axis lie at the end of umbilical spiral cords, the outermost of which is mostly submerged beneath a thick umbilical veneer. The third tooth is situated adjacent to the floor of the aper- Page 174 Figures 66, 65. Oblique spiral view. 68. Oblique basal view. Figures 67, 69-72. THE NAUTILUS, Vol. 121, No. 4 Tegula (s.1.) tridentata (Potiez and Michaud, 1838). UWBM 9789S, DV 1372-1, Recent, width = 9.9 mm. 66. Tegula (s.1.) melaleucos (Jonas, 1$44). 67. UWBM 97901, Paracas Hotel, Recent, oblique spire view, width = 23.9 mm. 69. OSU 37609, DV 211-3, late Pleistocene, lateral view, length = 28.2 mm. 70. UWBM 97901, oblique basal view. 71. OSU 37609, apertural view. 72. OSU 37609, oblique basal view, width = 29.0 mm. Figure 73. Tegula (s.1.) (?) rubroflammulata (Koch in Philippi, 1843). OSU 3 7610, DV 341, early Pleistocene, oblique basal view, width = 22.0 mm. Figures 74-78. Tegula (s.1.) masiasi new species. 74. UWBM 97904, DV 475-1, early Miocene, lateral view, length = 16.6 mm. 75. UWBM 97903, DV 1019-1 . holotype, middle Miocene, oblique spire view, width = 15.9 mm. 76. UWBM 97903, basal view. 77. UWBM 97905, DV 1648-1, early Miocene, lateral view, length = 14mm. 78. UWBM 97905, basal view showing edge of umbilical area, width = 16.9 mm. ture. The inside of the outer lip often has four to six short elongate teeth. In the latter three characters the species resembles Cantallocostoma quadricostatum, with which it has been grouped using mitochondrial DNA sequences by Hellberg (1998), who placed the species with Agathi- stoma. In its color and obsolete spiral sculpture, how- ever, “tridentata” specimens greatly resemble Peruvian species of Chlorostoma, to hich they were assigned by Marincovich (197: and Guzman et al, (1998). The only fossil example of T. tridentata in Peru comes from a middle neler. marine terrace bed near San Juan de Marcona, southern Peru. Tegula (s.1.) melaleucos (Jonas, 1544) (Figures 67, 69-72) Trochus melaleucos Jonas, 1544: 169; Philippi, 1846, Die kKrei- selschnecken oder Trochoideen, p. 185, pl. 28, fig. 16. Tegula melaleucos (Jonas). —Dall, 1909: 239. Tegula ( Agathistoma) melaleucos (Jonas, 1S44).—Keen, 1971: 340, fig. 106; Alamo and Valdivieso, 1997: 13, fig. 22. Not Tegula (Agathistoma) melaleucos (Jonas, 1S44).—DeVries 1986: 515, pl. 27, figs. 1, 2, 10, 12 [possibly Tegula (s.1.) rubroflammulata ( (Koch in ayer 1843) ]. Material Examined: OSU 37609, DV 211-3, late Pleistocene, L = 28.2, W = 29.0: UWBM 97901, Paracas Hotel, Recent, L 17.3, W 23.9: UWBM 97902, northern Peru, Recent, L = (11.5), W = 20.1. Specimens assigned to Tegula melaleucos by DeVries (1986) but more likely belonging to Tegula (s.1.) rubroflammulata; OSU: 37610, DV 341, early Pleistocene: L = 22.0, W = 22.0; OSU 37611, DV 341, L = (14.8), W = (18.0). Occurrence: Recent: northern Peru; rarely in south- ern Peru. T. DeVries, 2007 Remarks: Specimens of Tegula melaleucos are charac- terized by a strongly bicarinate periphery, a line of pro- tractive nodes between the senale ry and suture, and protractive brown stripes laterally and basally, where they spiral into an open umbilicus. A single spiral cord emerges from the umbilicus and is truncated by a ar umellar ridge that ends in a basally projecting tooth. second prominent tooth protrudes at the juncture of : columella with the floor of the aperture, and additional small teeth may occur along the edge of a beveled ledge that passes just inside the floor of the aperture. Impertectly preserved specimens of Tegula from the uppermost Pliocene / lower Pleistocene Mancora Ta- blazo of northern Peru (Figure 73; DeVries, 1986; 1988) have less impressed sutures and more convex profiles than typical specimens of T. melaleucos and nodes near the suture that are not protractive. These specimens are better referred to T. rubroflammulata, a Recent species that had been reported to range only as far south as Colombia (Keen, 1971). Tegula (s.1.) masiasi new species (Figures 74-78) Diagnosis: Spire whorls with broad spiral cords; um- bilicus narrow, open; shell lacking purple-black outer layer. Description: Shell conical, up to 16 mm wide. Spire angle about 70 degrees. Periphery at base; angular. Su- tures appressed to impressed. Protoconch anion: te- leoconch with at least four flat-sided to convex whorls. Axial sculpture absent: colabral growth lines strongly prosocline. Spiral sculpture absent or with several broad, low tee cords on spire. Outer shell layer lacking purple-black color. Base flattened to weakly convex, without visible spiral sculpture. Inner side of outer lip smooth. Umbilicus open. Columella thin, with at least one tooth at end of umbilical cord. Parietal flange barely overhanging umbilicus. Type Locality: km east of mouth of Quebrada Gramonal, on bluff over- looking road to Funda $ Santa Rosa (Figure 79). 14°45'50"S, 75°30'22"/W (Lomitas 1:100,000 quad- rangle). Lower middle Miocene. Type Material: }UWBM 97903, DV 1019-1. holotype, early middle Miocene, L = (7), W = 15.9. Other Material Examined: U\WBM 97904, DV 47S8- ie early Miocene, L = 16.6, W = (21.8); UWBM 97905, DV 1648-1, early Miocene, L = 14.0, W = 16.9: MUSM INV 143, DV 1648-1, L = 14.6, W = 19.8 Occurrence: Early to early middle Miocene: southern Peru. Etymology: Named in honor of Antonio Masias, pe- troleum geologist from Arequipa, Peru, who has pro- vided advice on Peruvian matters since we both attended Oregon State University in 1977-1978. DV 1019-1, Gramonal, about one-half oe vo =I S| a a] AS a de Ullujalla @ Locality-sample 0 5 km SCALE Contour interval is 100 m. 600 } eee Figure 79. Type locality (DV 1019-1) of Tegula (s.1.) masiasi new species. Remarks: Specimens of Tegula (s.|.) masiasi are the oldest known teguline »s from the Pisco Basin: they occur near the base of a depositional sequence that unconform- ably underlies the Pisco Formation near Cerros Colorado (Figure 78), which implies the lower portion of the up- permost Oligocene to lower middle Miocene Chilcatay Formation (De Vries, 1998), as well as a few tens of meters above the base of the Pisco Formation, which indicates an early middle Miocene age (DeVries and Schrader, 1997). Despite the poor state of preservation, it does appear that these early and middle Miocene specimens are not chlorostomines, which. first appear with their purple-black outer shell layer in beds at Alto Grande at about 9-10 Ma (Muizon and DeVries, 1955: DeVries and Schrader, 1997). DISCUSSION Fossils found since 1990 offer a new perspective on the origin of Tegulinae. Tegulines had been thought to have arisen during the early or middle Miocene (Hickman and McLean, 1990), based on knowledge of fossil taxa from Japan and California [e.g., Tegula thea Nomland, 1917 Page 176 THE NAUTILUS, Vol. 121, No. 4 and T. varistriata Nomland, 1917, Santa ae beds, California, upper Miocene (Nomland, 1917); T. dalli Ar- nold, 1907, and T. dalli var. inornata ‘ciold: ae To- panga Group, California, middle Miocene (Arnold, 1907; Yerkes and Campbell, 2005); T. dalli arnoldi Addicott, 1970, Olcese Sand, California, uppermost lower Miocene (Addicott, 1970; Sanchez and Prothero, 2003) ]. The dis- covery of T. jeanae Squires and Saul, 2005, a late Cam- panian species from the Chico Formation of California with many attributes of Chlorostoma (ventricose whorls, black-brown outer shell layer, absence of beaded spiral cords), recasts discussions of teguline phylogeny (Squires and Saul, 2005). A flat-sided, spirally bended Cretaceous teguline, the late Maastrichtian T. ovallei Philippi, 1887, from central Chile (Bandel and Stinnesbeck, 2000), fur- ther demonstrates ri pre-Miocene diversity and. geo- graphic distribution of tegulines. As a consequence of these Cretaceous discoveries, an evolutionary scenario rejected by Hickman and McLean (1990), which placed the plesiomorphic Tegula near the base of the trochid group, gains credence, whereas sce- narios inspired by the molecular data of Hellberg (1998) that eae a strictly late Neogene timeline for teg- uline phylogeny are undermined by these Cretaceous data, as sell as by early, middle, and late Miocene oc- currences of tegulines in Peru. The following observations provide some further con- straints on phy logene tic hypotheses related to Tegulinae. PERUVIAN CHLOROSTOMINE GROUP The Peruvian chlorostomine group is comprised of four modern species: Chlorostoma atrum, C. ignotum, C. eu- ryomphalum, and C. luctuosum. Cradatoas of sculpture indicate the latter two species may be one; C, luctuosum would be the senior synonym. Adding C. quipua extends the record of Peruvian ealonsctowines to 9 Ma (Muizon and DeVries, 1985). This late Miocene occurrence pre- cludes Hellberg’s (1998) molecularly based hypothesis that these chlorostomines appeared in western South America during the late Pliocene and constitute a sister group to a North Atlantic Pliocene group of Agathistoma Species. The flattened base, open umbilicus, and spiral threads of Chlorostoma quipua indicate that it is most closely related to the modern C. luctuosum. Some specimens of C. quipua from upper Pliocene beds above Playa Huacllaco also have broad spiral cords like those on some specimens of modern C. atrum. A single upper Pliocene Hu: 1c lle ico spe cime n hi iS strongly convex spire al cords like those on specimens of the modern Chilean C. ignotum. This morphological variation suggests the onset of a ra- diation of South American chlorostomines near the end of the Pliocene. The late Pliocene was also a time of species-level molluscan mass extinction throughout the Peruvian Faunal Province (DeVries, 2001) The origin of chlorostomines in Peru and Chile is not known. Tegulines are found, rarely, in lower and middle Miocene beds of southern Peru (T. (s.].) masiasi; Figures g 74-78), but they do not resemble late Neogene Peruvian Chlorostoma. Of Chilean Miocene species assigned to Chlorostoma (Nielsen et al., 2004), none edu the purple-black outer shell layer that characterizes the type species of Chlorostoma and most moder Peruvian chlo- rostomines. At three mm in length, Tegula austropacifica Nielsen, Frassinetti, and Bandel, 2004, is extraordinarily small for a chlorostomine. Tegula chilena Nielsen, Fras- sinetti, and Bandel, 2004, lacks the flattened base of chlorostomine species. Tegula matanzensis Nielsen, Frassinetti, and Bandel, 2004, resembles Californian specimens of C. funebralis, as the authors note, but the specimen lacks critical portions of the columella from which a better comparison can be made. Chlorostomines appeared in southern Peru at the same time as the turbinid genus, Prisogaster Mérch, 1850 (DeVries, 2006), and mollusks with Panamic affini- ties (DeVries, 2006), an immigration pattern consistent with either a western North American or boreal Asian origin for the group. Molecular data (Hellberg, 1998) ‘acest Peruvian chlorostomines are more similar to Californian than Asian taxa. Shell characters are equivo- cal on this point. Protractive stripes, present on speci- mens of the oldest Peruvian chlorostomine, C. quipua, are also seen on some specimens of Californian C. gallina, C. rugosum, and Asian C. nigerrimum and C. rusticum. Protractive ribs, which occur in rare examples of C. quipua, are found on some specimens of the Cali- fornian “Tegula” brunnea and several Asian species. Im- bricate subsutural spiral cords, which occur rarely on juvenile whorls on specimens of C. atrum, are most char- acteristic of the Californian C. fiumebralis (Figures 5, 6), are weakly developed on specimens of the Califor nian C. gallina and C. rugosum, and are not seen on specimens of Asian species. Peruvian chlorostomines are distinguished from all Californian chlorostomines and all Asian taxa except a few specimens of C. rusticum by possessing a very thin and expansive umbilical veneer ‘that does not bury the spiral umbilical cord. A thicker umbilical veneer di rapes across the spiral umbilical cords in Californian and Asian specimens, largely burying the spiral umbilical cord and leaving visible only a blunt adaxial columellar tooth, a tooth at the base of the columella, and an interve ning depressed nacreous wedge (e.g., C hlorostoma funebralis; Fie. 6). Peruvian Ghilbrestoutines also lack a second well developed tooth at the base of the columella, a character usually seen on Californian and Asian chlorostomine specimens (Fig. 6). These two derived characters—thin umbilical veneer, obsolete basal columellar tooth—-may indicate that Peruvian chlorostomines are a sister group to Californians Asian chlorostomines, with a common an- cestor in the North Pacific Ocean older than late Mi- ocene. CANTALLOCOSTOMA GROUP Cantallocostoma is an endemic western South American genus characterized by beaded spiral cords and two T. DeVries, 2007 Page 177 adaxially situated spiral umbilical cords. Cantallocostoma panistostum appears first in upper Miocene beds of southern Peru with other Panamic species (DeVries, 2002). It and the extant C. quadricostatum are unlike any Neogene or Recent teguline from Peru or Chile. Speci- mens of Cantallocostoma share with specimens of Inti- stoma the presence of two adaxially situated spiral um- bilical cords (mostly covered by a thick umbilical veneer in specimens of Intistoma) and an erect columella with little in the way of a parietal flange or callus. Genetic data of Hellberg ( (1998), however, show no close affinity be- tween C. quadricostatum and I. aureotinctum. INTISTOMA GROUP Intistoma has been created to include two very similar species, the modern Californian I. aureotinctum and early Pliocene Peruvian I. pirqua. Their distinctive spiral sculpture ( (strong bicarinate periphery, protractive sub- sutural nodes, thre -e broad basal spiral cords), absence of a purple-black outer shell layer, absence of apertural teeth, and the isolation of T. aureotinctum in mtDNA phylogenies (Hellberg, 1998) indicate that neither Chlo- rostoma nor Agathistoma properly encompass these taxa. Two other groups, one consisting of the modern north- ern Peruvian / Panamic Tegula (s.1.) melaleucos and pos- sibly T. (s.1.) rubroflammulata, the other comprising Mi- ocene and Pliocene species from California [e.g., T. (s.1.) dalli\, exhibit the distinctive subsutural spiral row of pro- tractive nodes of Intistoma, but both lack the three broad basal spiral cords that characterize the new genus. The Californian Neogene species also lack the differentiation of spiral sculpture (coarse primary spiral cords, fine ter- tiary spiral threads overrunning primary spiral cords and interspaces) that is visible on well-preserved specimens of I. aureotinctum. Tegula (s.l.) tridentata (Potiez and Michaud, 1838) Tegula (s.1.) tridentata is a small teguline with a record in Peru and Chile extending no farther back than the middle Pleistocene. The distinctive purple-black exterior is shared with Peruvian chlorostomines, but the arrange- ment and number of columellar teeth is like that of Panamic species of Agathistoma. Mitochondrial DNA data (Hellberg, 1998) are not helpful on this point of phylogeny, as T. (s.1.) tridentata usually clusters with Cantallocostoma quadricostatum, which it resembles in only one significant shell character: two adaxial umbilical spiral cords, both terminating in a columellar tooth. For now, the proper phylogenetic assignment of T. (s.].) tri- dentata remains elusive. AUSTRAL AGATHISTOMA Panamic agathistomines, which are so speciose in warm waters of the Panamic Faunal Province, have been no- tably unsuccessful in penetrating the cold waters of the Peruvian Faunal Province. The only fossil agathostomine from western South America is a specimen from upper Pliocene beds of southern Peru assigned to Agathistoma patagonicum. A Miocene agathistomine reported from central Chile, Agathistoma antiquum Nie Isen, Fras- sinetti, and Bandel, 2004, a Miocene occurrence of A. patagonicum reported from Argentina (Ihering, 1907; del Rio, 1998), and an extensive record of modern Aga- thistoma patagonicum from southern Brazil to southern Argentina, including the Magellanic waters of Argentina (Carcelles and Williamson, 1951), if the synonymy of A. fuscesens and A. orbignyana with A. patagonicum is ac- acer (Forcelli, 2000), may point to a Miocene austral origin for the subgenus ( Nielsen et al., 2004), rather than the Pliocene western Atlantic origin suggested by Hick- man and McLean (1990). ), Alternatively, the monophy ly of Agathistoma might be suspect; compact, beaded, rule toothed austral tegulines with or without open um- bilici might constitute a long-lived sister group to a group of Caribbean and Central American species. CONCLUSIONS Pre-late Miocene tegulines in southern Peru are exceed- ingly rare, poorly preserved, and bear little resemblance to late Neogene or extant taxa. During the early late Miocene, at least two lineages of tegulines, Chlorostoma and Cantallocostoma, entered Peruvian waters. [t is un- clear whether these two genera originated in California or Asia. Other trochids, namely Diloma Philippi, 1845, are thought to have dispe srsed across the Equator and across fie Pacific Ocean from Australia, rafted by buoy- ant fragments of the brown kelp, Durvillaea Bory de Saint-Vincent, 1826 (Donald et al., 2005). lies of Pe- ruvian Tegula likewise live upon on brown kelp (Lessonia Bory de Saint-Vincent, 1825) (Véliz and Vasquez, 2000; V. Mogollon, pers. comm., 2006), and thus may have been rafted to western South America from California or Asia in the same manner as Diloma. Species of Chlorostoma and Cantallocostoma re- mained relatively unchanged until the end of the Pliocene, when a mass extinction swept away 50 percent of molluscan species in the Peruvian Faunal Province (DeVries, 2001). At that time, chlorostomines experi- enced a mini-radiation in southern Peru or Chile, with one species, C. atrum, eventually spreading to southern Argentina (Carcelles and Williamson, 1951). Cantallo- costoma panistostum was replaced at the same time by the modern C. quadricostatum. Another lineage of tegulines, represented by the early Pliocene Intistoma pirqua, appeared on Peruvian shores by the early Pliocene. Although the genus is now extinct in Peru, it persists in California in the guise of I. au- reotinctum. It is likely that these species, with their br« vad basal spiral cords, are not related to the “dall” lineage of Californian tegulines that ranged from the early Miocene to Pliocene (Addicott, 1970). Page 178 THE NAUTILUS, Vol. 121, No. 4 ACKNOWLEDGMENTS I would like to thank G. Collado (Facultad de Ciencias, Universidad de Chile), L. Groves and J. McLean (Natu- ral History Museum of Los Angeles County, USA), S. Rugh (San Diego Museum of Natural History, USA), J. Vasquez Umveradad Catolica del Norte, Coquimbo, Chile), D. Véliz (Universidad de Chile, Santiago), G. Pas- torino (Museo Argentino de Ciencias Naturales, Buenos Aires), S. V. Mogollon ( (Universidad Nacional Federico Villarreal, Lima, Peru), and W. J. Zinsmeister (Purdue University, West Lafayette, Indiana, USA) for access to museum collections, loans of specimens, or valuable ad- vice. Helpful critiques were offered by D. Zelaya and M. Griffin (Museo de Ciencias Naturales, La Plata, Ar gen- tina). Support for field research was provided in part by a Fulbright scholarship in 1999. LITERATURE CITED Adams, A. 1853. Contributions towards a monograph of the Trochidae., a family of gasteropodous mollusks. 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Wood, W. 1828. a Seca to the Index Testaceologicus; or, a catalogue of shells, British and foreign, arranged accord- ing to the Linnean system. London, 59 pp- Yerkes, R. F. and R. H. Campbell. 2005. Preliminary Geologic Map of the Los Angeles 301 x 601 Quadrangle, Southern California. United States Geological Survey, Open-File Report 2005-1019. Locality-samples. GPS = latitude and longitude coordinates obtained by a hand-held GPS unit. GE = coordinates obtained from satellite images available on-line from Google DV 211-3 DV 341-1 DV 381-5 quad angle). Middle Pleistocene, DV 382-1 75°05'26"W ( DV 3958-1 DV 401-1 Recent. DV 463-1 DV 470-1 Lower terrace, five km north of Chala (Chala 1:100,000 quadrangle). Above ee road from Hacienda Tunca to Quebrada Huaricangana; hillside of brown sandstone. Earth™., Punta Lobitos, northern Peru, western point, terrace surface midway between sea cliff and inshore edge of deposit. 04°27'12"S, S1°17'25"W (GE; Lobitos 1:100,000 quadrangle). Quebrada Mogollon, northern Peru. Upper coquina of Mancora Tablazo. Lower Pleistocene. San Juan- Lomas road, km 47.5. Uppermost coquina. 15°22'59"S, 75°03" 11"W ( Upper Pleistocene. San Juan 1:100,000 San Juan / Lomas road, kilometer marker 50, flat-topped knoll south : highway. 15°22'02"S, San Juan 1:100,000 quadrangle). Remnant of marine terrace. Playa Canastones, Bahia de la Independencia, Peru (Punta Grande 1:100,000 quadrangle). Recent. Hueco La Zorra, north end of beach. 14°02'31"S, 76°15'51"W (Punta Grande 1:100,000 quadrangle). Upper Pleistocene. Upper Pleistocene. 14°56’ S. 75°09’ W (GE: Palpa 1:100,000 quadrangle). Lower Pliocene. DV 478-2 Lomas : hileatay, northeast end of outcrop. 14°11'42"S 76°06'57"W (Punta Grande 1:100,000 quad- rangle). Chile ‘atay Formation, lower Miocene. DV 571-1 Alto Grande, about one km south of intersection with abandoned paved road to San Juan de Marcona, on south-facing hillside west of Panamerican Highway; one of several shell banks. 15°26'57’S, 74°52'06"W (Acari 1: 100,000 quadrangle), Yauca, roadcut on western side of Panamerican Highway as it descends to valley floor. 15°39'49"S, 74°31'50"W (Yauca 1:100,000 quadrangle). Camu about one-half km east of canyon mouth. 14°45'50"S, 75°30'22"W DV S09-1 DV 1019-1 quadrangle). Middle Miocene. DV 1029-1] quadrangle). Lower Pliocene. DV 1032-2 ). Middle upper Miocene. Pisco formation. Lower Pliocene. (Lomitas 1: 100,000 Yauca Depression, west of Pana merican Highway. 15°39'29"S, 75°35'08"W (GPS, Yauca 1:100,000 Morro Abra de los Chaparrinos, descending from highest terrace level, north and south of second curve in Panamerican Highway. 15°52'59"S, 74°10'05"W (Chala 1:100,000 quadrangle). Upper Pliocene. DV 1252-1 above non-marine de posits. 15°45'56"S, DV 1254-Bal 6 rangle). Upper lower Pliocene. DV 1254-Bal 10 Section along Panamerican Highway, meters above basement rocks in measured section. 15°53'25"S quadrangle). Upper Pliocene. Quebrada de la Vaca, roadcut along Pa inamerican Highway, south of south wall, uppermost terrace 74°18'50"W (GPS; Chala 1:100,000 quadrangle). Section along Panamerican Highway, ten km southeast of Chala and above Playa Huaclla ico. 35 meters above basement rocks in measured section. 15°53/25"S, 74°09'52"W (GPS; Chala 1:100,000 quad- ten km southeast of Chala and above Playa Huacllaco, 47.5 ea 95"5 — 74°09'52"W (GPS: Chala 1:100,000 DV 1254-] Sacaco, shell banks southwest of north-south road to farmhouse (chacra). 15°33'03 S', 74°43'50"W (GE; Yauca 1:100,000 quadrangle). Lower Pliocene. DV 1355-1] Quebrada Pongo, one km upstream _ juncture with Quebrada Caracoles. 15°30'22"S, 74°45'40"W (GPS; Yauca 1:100,000 qui adrangle) Upper Pliocene. DV 1372-1] tocky beach on northwestern side ge Punta Lomas (Acari 1:L00,000 quadrangle). Recent. DV 1418-1] East side of Acari Depression, 15°34'50"S, 74° 36'59°W (GPS; Yauca 1:100,000 quadrangle). Upper Pliocene T. DeVries, 2007 Page 15] DV 1598-1 DV 1599-1 DV 1635-1 DV 1648-1 DV 1713-1 JM 82-19 JM 82-20 WIZ 345 Isla Ipun Paracas Hotel Roadcut oO Panamerican Highway, descent from north into Yauca. Shell beds. 15°39'49"S, 74°31'50"W (GPS: Yauca 1:100,000 quadrangle). Lower Pliocene. ae La Zorra, north end of beach (see DV 401-1). Recent. Yauca re aes west of Panamerican Highway. 15°39'33"S, 75°34/54"W (GPS; Yauca 1:100,000 quadrangle). Lower Plioce ne. Westward- facing side of valley, southwest of Cerros Colorado. 14°22'25"S, 75°53'52”/W (GPS; Punta Grande 1:100,000 quadrangle). Marine terrace on east side of Panamerican Highway north of road to Acari. 15°36'09"S, 74°41'08’"W (GPS; Yauca 1:100,000 quadrangle). Lower Pleistocene. Cerro El Huevo, northeast of San Juan de Marcona. 15°18’ S, 75°09’ W (San Juan 1:100,000 quadrangle). Upper Pleistocene. Cerro E] Huevo, northeast of San Juan de Marcona. 15°18’ S, 75°09" W (San Juan 1:100,000 quadrangle). Upper Pleistocene. Coquimbo, Chile. Pleistocene. Approximately 29°58" S, 71°20' W (GE). Isla Ipun, Chile, shores of eastern embayments. 44°38’ S, 74°44’ W (GE). Recent. Beach south of Hotel Paracas, facing Bahia Paracas, southern Peru. 13°50'09"S, 76°15’ 19”"W (GE, Pisco 1:100,000 quadrangle). THE NAUTILUS 121(4):182-190, 2007 Page 182 Three new species of Paryphantopsis (Gastropoda: Pulmonata: Charopidae) from the Nakanai Mountains, New Britain, Papua New Guinea John Slapcinsky Robert Lasley Florida Museum of Natural History and Department of Zoology University of Florida Gainesville, FL 32611 USA slapcin@flmnh.ufl.edu ABSTRACT Recent surveys (February-March, 2005) of the terrestrial snail fauna of the Nakanai Mountains, central New Britain, Bismarck Archipelago have uncovered several undescribed species, in- cluding three new species of Paryphantopsis, a diverse genus of charopid snails, previously believed to be endemic to mainland New Guinea and adjacent islands of the Louisiade Archipelago. The three species are described using shell, genital, and udu ww morphology. Although the land snail fauna of New Britain is arguably the best sampled in Papua New Guinea, there has been little sz unpling in the interior mountains of the Nakanai, Whiteman, Baining, and Willaumez ranges, which harbor pre- viously undetected species of terrestrial snails. Additional Keywords: Pulmonata, Charopidae, Paryphantopsis, New Britain, Papua New Guinea INTRODUCTION This is the third in a series of reports on the results of recent field surveys for terrestrial mollusks from Papua New Guinea. The previous two reports focused on spe- cies of Paryphantopsis from the mainland New Guinea (Sk apcinsky, 2005) and nearby is- lands of the Louisiade Archipelago (Slapcinsky, 2006). This report reviews Paryj shantopsis species collected during three weeks of field surveys in February and March 2005 from the N Pomio, East New Britain Province, Papua New Guinea Figure 1). Paryphantopsis, a diverse genus of charopid snails endemic to New Guinea and surrounding islands, is comprised of 23 described species | (Solem, 1970: Slape insky, 2005: Sl ape ‘insky, 2006) that are distributed from Papua (Irian Jaya) to the Louisiade Archipe lago and that, prior to this ‘study, were not known from New Britain Nearly all species of Paryphantopsis have distributions stricted to single mountain ranges or islands where eastern peninsula of Nakanai Mountains northwest of they are found in hilly or mountainous terrain from 60 to 4000 meters elevation. In upland habitats they can be the most abundant snail species reaching densities of more than 10 individuals per square meter on Sudest and Ros- sel Islands in the Louisiade Archipelago (personal obser- vation). Despite their small size (4-12 mm), their bright yellow body coloration and ia ul activity in exposed locations on tree trunks, vegetation and rotting wood make them among the most hanes snail species. These showy snails are under- sampled and recent surveys have near ly doubled the number of recognized species. Fur- ther sampling in New Guinea and surrounding islands will certainly uncover additional species of Pary) phantop- sis as well as many other more cryptic snail species. New Britain, a large volcanic island, 35145 km?, ex- tending from 148° to 152° E longitude and from 4° to 7° S latitude, emerged in the late Miocene ( S—LO mya). The island’s basement rocks were deposited by volcanic ac- tivity, between the upper Eocene and the middle Oli- gocene. Volcanic activity ceased in the early Miocene le ading to regional subsidence and the de “position of ex- tensive reef comple xes. At the end of the Miocene, sub- duction of the Solomon plate under the Bismarck plate led to renewed volcanism, resulting in a chain of young arc volcanoes along the northwestern coast of New Brit- ain (Woodhead et al. 1998). At the same time, reef com- plexes along the south and eastern coasts of the island were apidly uplifted, creating extensive karst mountains. One of these ranges, the Nakanai Mountains, contains the Southern He ‘misphe re’s deepest caves, some nearly 1200 meters in depth (Audra et al. 2001). New Britain's isolation, complex geology, and extensive raised lime- stone have allowed the radiation of a diverse and 1: urgely endemic land snail fauna (Rensch, 1934; 1937 ) that is among the best surveye din Papua New Guinea. How- ever, rough karst terrain and lack of roads have limited nearly all surveys to coastal lowlands, and most taxa from Nakanai, Whiteman, Baining, and Willaumez ranges are still poorly surveyed (Beehler, interior mountains of the J. Slapcinsky and R. Lasley, 2007 Page 183 151.4°E 151.6°E Sy <~.... Nakanai 2 Mountains | PRS) ) Sas Ss } / A ge TRS \ / LASSE \ Jacquinot Bay on Figure 1. Distribution of Paryphantopsis in New Britain, Papua New Guinea; 0 = other sites sampled. 1993), even for macrofauna such as mammals (Flannary, pilasters, PR = penial retractor muscle, SD = spermathecal duct, SP = spermatheca, VA = vagina, VD = vas deferens. Terminology of vegetation types follows Paijmans (1976). Specimens are deposited in the following institu- tions: Bernice P. Bishop Museum, Honolulu (BPBM); Florida Museum of Natural History, Gainesville (UF); Natur-Museum Senckenberg, Frankfurt (SMF): Papua New Guinea National Museum, Port Moresby (PNGNM); Wroclaw University Museum of Natural His- tory (MNHW),. SYSTEMATICS Family Charopidae Hutton, 1884 Genus Paryphantopsis Thiele, 1928 Type Species: Flammulina (Paryphantopsis) lamel- ligera Thiele, 1928, by original designation. Paryphantopsis corolla new species (Figures 2-7, Table 1) 7 ; : ; ae ; Holotype: UF 366508, J. Slapcinsky, 25 February 1995: 12) and birds (Orenstein, 1976). The invertebrate : YP . J I : 3 / : icon 2 Negi . elie 2005. fauna of New Britain’s interior mountains is almost en- tirely unknown. This is disturbing because intensifying Paratype: UF 366453 (1 specimen), J. Slapcinsky, 25 land usage, including logging and the establishment of oil palm plantations (McAlpine and Fryne, 2001), threatens to deforest extensive areas of New Britain before they can be adequately inventoried, potentially leading to largely undocumented losses in biodiversity. MATERIALS AND METHODS Specimens were hand-collected, drowned overnight, and then preserved in 75% ethanol. Gross anatomical dissec- tions were made under 75% ethanol using a dissecting microscope. Radulae were isolated from dissected buccal masses using 5% sodium hypochlorite solution. Scanning electron micrographs of radulae were made using a Field Emission SEM. Line drawings of the genital anatomy were made from digital images, and measurements were taken using an ocular micrometer. Shell measurements were made as figured in Slapcinsky (2005). The following abbreviations are used in figures of genital anatomy: AT = atrium, DI = diverticulum, EP = epiphallus, OV = free oviduct, PE = penis, PG = prostate gland, PP = penial February 2005. Type Locality: Papua New Guinea, East New Britain Province, New Britain, 12 km northwest of Marmar Village on the trail to Pakia Village, 5.432° S, 151.460° E, 900 meters altitude (Figure 1). Habitat: Collected in mixed hill forest on vegetation within 1 m of the ground. Description: The adult shell is depressed-globose, small for the genus, 3.5-3.6 mm (mean = 3.55, see Table | for sample size and standard deviation) in diam- eter and 2.2-2.3 mm (mean = 2.25) in height, with 2.5- 2.6 (mean = 2.55) rapidly expanding whorls (Fig- ures 24). The apical surface of the whorls flattens be- tween the deep suture and the shell periphery which is slightly angular a little above its mid-point. The basal surface of the whorl is evenly rounded from the angular periphery to the umbilicus. The spire is slightly elevated, 0.1—0.2 mm (mean = 0.15). Teleoconch whorls descend slowly and regularly and shell height/diameter ratio is 0.61-0.66 (mean = 0.63). Approximately 1.3 rounded protoconch whorls are sculptured with L3 rows of spiral Table 1. Measurements in mm of undamaged adult shells of three species of Paryphantopsis. Species N H D SH SD AH AW W P. corolla 2 Mean+=SD 2.25+0.07 355+0.07 0.15+0.07 1604000 160+000 215+007 2.55 +0.07 Range 2.9-2.3 3.5-3.6 0.1-0.2 1.6-1.6 1.6-1.6 2.1-2.2 2.5-2.6 P. fragilicosta 6 Mean+SD 3.43+018 5.00+020 0.134005 2334012 248+016 292+0.16 3.00+0.09 : Range 3.2-3.7 4,8-5.3 0.1-0.2 2.9-9.4 2.3-2.7 2.83.2 2.9-3.] P. nucella 10 Mean+SD 3.75+0.32 5.22+0.23 0.18+0.04 2.06+020 2.784027 316+0.22 281+0.12 Range 3.34.1 4.9-5.7 0.1-0.2 1.8-2.5 2.4-3.3 2.9-3.6 2.6-3.0 N = number of specimens, H = height, D = diameter, SH = spire height, SD = spire diameter, AH = aperture height AW = aperture width. W = number of whorls Page 184 THE NAUTILUS, Vol. 121, No. 4 a! Sa Figures 2-7. Paryphantopsis corolla, 2-4. Shell, Holotype UF 366508, Scale bar = 1 mm; Figure 5, Genitalia, UF 366453. Scale bar = 1 mm. 6-7. SEMs of radula, UF 366453. 6. Central and lateral teeth. 7. Marginal teeth. Scale bars = 10 xm, mss J. Slapcinsky and R. Lasley, 2007 Page 185 pits, which continue on the teleoconch where the pits elongate nearly fusing to form incised spiral lines. These apical pits are typical of nearly all species of Paryphan- topsis as, for example, in Pary) shantopsis louisiadarum (see Solem, 1959: plate 13, fig 6). Approximately every third growth line is accentuated with a pe sriostracal ex- tension that bears a sharply pointed triangular process at the shell periphery. These processes are approximately 0.2 mm in length and 0.3 as wide at the base. The pro- toconch and teleoconch whorls are uni formly brown ex- cept for the columellar edge of the peristome and um- bilicus, which are slightly darker brown. A reflection of the peristome covers approximately 0.4 of the umbilicus. The aperture is large and ovate with an aperture:width to aperture:height ratio of 1.31-1.35 mm (mean = 1.34). The external body color is bright yellow in life and there are no bands or other color patterns. Specimens preserved in ethanol fade to uniform cream. The head is short similar to other species of Paryphantopsis and the posterior of the foot is slightly shorter than average for the genus. The vas deferens is 0.2 the diameter of the head of the epiphallus which bears a short subapical diverticulum approximately 0.1 the length, and 0.6 the diameter of the epiphallus where they join (Figure 5 the epiphallus is uniform in diame the basal 0.3 is twice as wide and ovoid. The penial retractor muscle is long and originates at the diaphragm, inserting 0.6 the way to the base of the epiphallus. The penis is approxi- mately the same length as the epiphallus and the same diameter as the apical 0.7 of the epiphallus. The penis, narrow apically, widens slightly below the apex, and tapers basally to 0.2 the diame of the atrium where they join. The atrium, widest at the insertion of the penis, narrows abruptly by 0.6 at the gonopore. The vagina narrows slightly at the junction with the free oviduct and spermathecal duct. The S-sh iaped free oviduct is 1.2 times the width of the spermathecal duct at their junc- tion with the vagina. The diameter of the spermathecal sea doubles from its junction with the vagina distally to 0.4 the length, then narrows to 0.4 its greatest diameter at 0.7 the length, and remains narrow to the junction with the ovate spermatheca. The central teeth of the radula (Figure 6, second from the right) are tricuspid, 7-S zm wide and 11-12 pm long, of similar shape and length but slightly narrower and shorter than the first laterals, 8-9 fxm wide and 12— 13 pm long (Figure 6). The mesocones of both the cen- trals and first laterals are tall and slender, and project beyond their basal plates. The ectocones of the central teeth are trigonal and symmetric. Ectocones and en- docones of the laterals are trigonal and about 0.5 the height of the mesocones. The endocones of the laterals are slightly larger but otherwise of similar shape to their ectocones. The first 5 teeth to the left and right of the central row are similar to the first lateral teeth, the next 3 teeth on either side grade in shape and are difficult to classify as either lateral or marginal teeth. The last 4 marginal teeth are wider, 10-11 pm wide, and shorter ). The apical 0.7 of 7-S pm long (Figure 7). Both the endocones and ecto- cones of the marginal teeth are irregularly multicuspid. The endocones are 0.9-0.8 as tall as the mesocones and usually bear 3 cusps, although these are sometimes divided into additional cusps. The ectocones sit on a rectangular base and are divided into 5 or more cusps that are 0.7 to 0.8 the height of the mesocones. Remarks: The only other Paryphantopsis species with shells bearing periostracal processes are: P. abstrusa Slapcinsky, 2005; P. fultoni (Coen, 1922); P. lebasii Slapcinsky, 2005; P. lame llizera (Thiele, 1928): P. striata (Fulton, 1902); P. yawii Slapeinsky, 2005; and P. yelensis Slapcinsky, 2006. However, the periostracal processes in P. corolla do not overlap unlike P. fultoni and P. yawii, and are sharply pointed unlike P. lebasii. Also, the pro- cesses are prominent unlike P. abstrusa and P. yelensis. Finally, the shells of P. lamelligera and P. striata are more than twice the size of P. corolla. Paryphantopsis corolla has an epiphallus that is basally robust unlike all other Paryphantopsis species for which the genital anatomy is known e xcept for P. nucella and P. misimensis Slapcinsky, 2006. In contrast to P. corolla, P. nucella has an apically inflated penis in addition to a large dark glan- dular area on the spermathecal duct, and P. misimensis lacks an apical diverticulum. The ectocones of the mar- ginal teeth sit on a rectangular extension of the tooth, unlike all other Paryphantopsis tor which the radular morphology is known other than P. fragilicosta and P. nucella. Etymology: The species name is from the Latin femi- nine noun corolla and describes the shells resemblance to a small garland or crown. Paryphantopsis. aa new species (Figures 8-13, Table 1 Holotype: UF 353995, J. Slapcinsky, 2S February 2005. Paratypes: Papua New Guinea, East New Britain Province, type locality, J. ae 28 February 2005, BPBM 282461 (2 specimens), PNGNM (1 spe cimen), UF 366505 (4 specimens), UF 366507 (9 specimens). Type Locality: Papua New Guinea, East New Britain Province, New Britain, Saukale, 13 km northwest of Marmar Village on the trail to Pakia Village, 5.426° S, 151.453° E, 910 meters altitude (Figure 1). Habitat: Collected in mixed hill forest on vegetation within | meter of the ground. Description: The adult shell is globose to depressed- globose, average size for the genus, 4.5-5.3 mm (mean = 5.00, see Table 1 for sample size and standard deviation) in diameter and 3.2-3.7 mm (mean = 3.43) in height, with 2.9-3.1 (mean = 3.00) rapidly expanding whorls (Figures 8-10). The suture is deeply impressed and the shell periphery is evenly rounded. The spire is slightly elevated, 0.1—-0.2 mm (mean = 0.13). Teleoconch whorls descend slowly and regularly until the end of the body whorl, which desce its. slightly more rapidly. The shell Page 186 THE NAUTILUS, Vol. 121, No. 4 aes Figures 8-13. Paryphantopsis fragilicosta. 8-10. Shell, Holotype UF 383995. Scale bar = 1 mm. 11. Genitalia, UF 366507. Scale bar = | mm. 12-13. SEMs of radula, UF 366507. 12. Central and lateral teeth. 13. Marginal teeth, Scale bars = 10 jm. J. Slapcinsky and R. Lasley, 2007 Page 187 height:diameter ratio is 0.65-0.71 (mean = 0.69). There are eee 1.4 evenly rounded protoconch whorls sculptured with 13 spiral rows of small pits which con- tinue on the teleoconch where the pits elongate nearly fusing to form discontinuous spiral striae. Short peri- ostracal extensions that do not bear periostracal pro- cesses are present approximately every four growth lines, These extensions are often worn or absent in older adult shells. The protoconch and teleoconch whorls are usually uniformly brown, shiny, and translucent, although older worn shells can be dull brown, opaque, with a white protoconch. The umbilicus is closed or nearly closed by a reflection of the peristome. The aperture is large and ovate, with an aperture:width to aperture:height ratio of 1.0S—1.28 (mean = 1.18). The external body color is bright yellow in life and there are no bands or other color patterns. Specimens preserved in ethanol fade to uniform cream. The head is short similar to other species of Paryphantopsis and the posterior of the foot is of average length for the genus. The vas deferens is 0.2 the diameter of the slightly inflated head of the epiphallus which does not bear a diverticulum (Figure 11). The epiphallus is approxi- mately 1.5 the length and 0.5 the diameter of the penis and widens only slightly at the junction with the penis. The penial retractor muscle is moderate in length, origi- nating from the diaphragm and inserting at the basal 0.3 of the epiphallus. The penis is robust and of uniform width apically, narrowing basally to 1.2 the width of the atrium just before their junction. The atrium narrows slightly and then broadens between the vagina and gono- pore. The vagina broadens slightly at its junction with the free oviduct and spermathecal duct. The straight free oviduct is 0.5 the width of the spermathecal duct at their junction with the vagina. The sper mathecal duct is rela- tively wide basally, tapering to 0.3 its basal diameter at its midpoint, and remaining narrow until the junction with the ovate spermatheca. The central teeth of the radula (Figure 12, middle row) are — S-9 pm wide, and 12-13 pm long, of similar shape and length but slightly narrower and shorter than the first lateral teeth, 10-11 xm wide and 14-15 pm long (Figure 12). The mesocones of the cen- tral and first later sal teeth are tall, slender, and project slightly beyond their basal plates. The ectocones of the central teeth are trigonal and symmetric. Ectocones and endocones of the lateral teeth are trigonal and about 0.5 the height of the mesocones. The endocones of the lat- eral teeth are slightly larger but otherwise of similar shape to their ectocones. The first 10 teeth to the left and right of the central row are similar to the first laterals. The next 3 on either side grade in shape and are difficult to classify as either lateral or marginal teeth. The last 4 marginal teeth are slightly wider, alot 11-12 pm wide, and shorter, 7-10 ym long (Figure 13). The base of the unicuspid or bicuspid endocones of the first and second marginal teeth is reduced, and they originate from the side of their mesocones. The ectocones are reduced, and their rectangular bases are usually devoid of cusps. The third and fourth marginal teeth often lack all cusps and are reduced to rectangular bases. Remarks: ~The only other Paryphantopsis species with shells that have periostracal extensions on the growth lines and no processes at the margin are: P. arcuata Jut- ting, 1964; P. dauloensis Solem, 1970; P. filosa Jutting, 1964: P. koragae Slapcinsky, 2005; P. latior Jutting, 1964; P. matawanensis Slapcinsky, 2005; P. platycephala Jut- ting, 1964; and P. pygmaea (Bavay, 1908). Paryphantop- sis fragilicosta is larger than P. filosa, is less depressed than P. dauloensis, has a higher spire than P. latior and P. platycephala, and has stronger spiral sculpture than P. pygiaca and P. arcuata. Paryphantopsis fragilicosta lacks an apical diverticulum, unlike all Paryphantopsis species for which the genital anatomy is known, except for P. louisiadarum, P. misimensis, and P. vanatinensis, which are much larger, as well as P. lebasii and P. yawii, which have long periostracal processes. The ectocones of the marginal teeth of P. fragilicosta sit on a rectangular extension of the tooth, unlike all other Paryphantopsis for which the radular morphology is known other than P. corolla and P. nucella. It differs from these species by having very few or no cusps on the marginal teeth. Etymology: The species name derives from the com- bination of the Latin adjective fragilis meaning crackling or easily a and the feminine noun costa meaning rib, and refers to the shells fragile periostracal exten- sions. Paryphantopsis nucella new species (Figures 14-19, Table 1) Holotype: UF 3583996, J. Slapcinsky, 258 February 2005. Paratypes: Papua New Guinea, East New Britain Province, New Britain: BPBM 282462 (2 specimens), MNHW MP LOOL (2 ease PNGNM (2 speci- mens), SME 329401 (2 2 specime ns) UF 383993 (54 speci- mens), UF 383994 (3 specimens), type locality, J. clea 28 February, 2005; UF 366504 (5 speci- mens), UF 366506 (7 specimens), 12 km northwest of Marmar Village on the trail to Pakia Village, 5.432° S, 151.460° E, 900 meters altitude, jes lapcinsky, 25 Feb- ruary 2005 Type Loc: ality: Papua New Guinea, East New Britain Province, New Britain, Saukale, 13 km northwest of Marmar Village on trail to Pakia Village, 5.426° S, 151.453° E, 910 meters altitude (Figure 1). Habitat: Collected in mixed hill forest on vegetation within 1 meter of the ground. Description: The adult shell is globose to depressed- globose, average for the genus, 4.9-5.7 mm (mean = 5.22, see Table | for sample size and standard deviation) in diameter and 3.1—-4.1 mm (mean = 3.75) in height with 2.6-2.9 (mean = 2.81) rapidly expanding whorls (Figures 14-16). The apical surface of the shell whorls Page 185 THE NAUTILUS, Vol. 121, No. 4 Figures 14-19. Paryphantopsis nucella. 14-16. Shell, Holotype UF 353996, Scale bar = 1 mm. 17. Genitalia, UF 383993 Scale bar | mm. 18-19. SEMS of radula, UF 383993. 18. Central and lateral teeth. 19. Marginal teeth. Scale bars = 10 wm J. Slapeinsky and R. Lasley, 2007 Page 189 are flattened between the deeply impressed sutures and the submedian shell periphery, which gives the shell a somewhat angular appearance. The spire is slightly elevated, 0.1-0.2 mm (mean = 0.18). The teleoconch whorls descend slowly and regularly until the end of the body whorl and more rapidly near the aperture. Shell height: diameter ratio is 0.62—0.78 (mean = 0.72). There are 1.5 evenly rounded protoconch whorls sculptured with spiral rows of small pits. Approximately 14 rows can be seen on the apex of shells. These pits become elon- gated on the teleoconch whorls nearly fusing to form weak incised spiral striae. Spiral striae w veaken, becoming obsolete on the final 0.30 of the body whorl where shell sculpture becomes predominated by growth lines all of which are accentuated with short periostracal extensions that do not bear peripheral processes. The protoconch is white, and teleoconch whorls gradually darken from yel- low brown to brown. The umbilicus is closed bya reflec- tion of the peristome. The aperture is ovate except for the flattened apical surface and has an aperture:width to aperture:height ratio of 0.94-1.25 (mean = 1.14). The external body color is bright yellow in life and there are no bands or other color patterns. Specimens preserved in ethanol fade to uniform cream. The head is short similar to other species of Paryphantopsis and the posterior of the foot is slightly shorter than average for the genus. The vas deferens is 0.2 the diameter of the head of the epiphallus, which bears an apical diverticulum that is approximately 0.3 the length and 0.7 the diameter of the midpoint of the epiphallus (Figure 17). The apical 0.7 of the epiphallus is uniform in diameter: the basal 0.3 is ovoid and twice as wide. The penial retractor muscle is moderate in length originating from the diaphragm and inserting 0.6 the way to the base of the epiphallus. The penis is 0.7 the length of the epiphallus. Its apical 0.3 is as robust as the base of the epiphallus and narrows rap- idly basally. The basal 0.7 of the penis is 0.3 narrower than the apex and uniform in diameter to the junction with the atrium. The atrium is 1.5 times wider than the penis at their junction and remains constant in diameter to the gonopore. The vagina expands slightly at the junc- tion with the free oviduct and spermathecal duct. The free oviduct is broad, not folded, and approximately the same diameter as the base of the spermathecal duct. The basal 0.3 of the spermathecal duct is wide and sur- rounded by darkly pigmented glandular tissue. The basal 0.5 of the spermathecal duct tapers apically to 0.2 of its original diameter and remains narrow until the junction mali the ovate spermatheca. The central teeth of the radula (Figure 18, middle row) are tricuspid, 10-11 zm wide and 14-15 pm long, and of similar shape and length, but are slightly narrower and shorter than the first lateral teeth, 11-12 wm wide and 16-17 pm long (Figure 18). The mesocones of the central and first lateral teeth are tall and slender, and project beyond their basal plates. The ectocones of the central teeth are trigonal and symmetrical. Ectocones and endocones of the lateral teeth are trigonal and about 0.5 the height of the mesocones. The endocones of the lateral teeth are slightly larger but otherwise of similar shape to their ectocones. The first 12 teeth on either side of the central row are similar to the first laterals. The next three on either side grade in shape and are difficult to classify as either iaeieale or marginals. The last four teeth are clearly marginals and are irregularly multicuspid, wider, about 13-14 pm wide, and shorter, 8—9 wm long (Figure 19). The endocones are as tall as the mesocones and are usually unicuspid, but sometimes bear 2 or 3 cusps. The ectocones sit on a rectangular base and are usually divided into 2-6 cusps that are 0.6 to 0.7 the height of the mesocones. Remarks: = Paryphantopsis nucella is similar to only P. koragae in having short periostracal extensions on all growth lines; ese growth lines do not bear processes at the shell periphery. The shell of P. koragae, however, differs in having the shell periphery above, rather than below, the midpoint of the whorl. Paryphantopsis nucella is unique among Paryphantopsis, for which the genital anatomy is known, in having a large glandular area on the base of the spermatheca and havi ing both an apically robust penis and a basally robust epiphallus. The ecto- cones of the marginal teeth sit on a rectangular extension of the tooth, unlike all other Paryphantopsis for which the radular morphology is known other than P. corolla and P. fragilicosta. Etymology: The species name derives from the Latin feminine noun nucella and describes the shells resem- blance to a small nut. Discussion and Conclusions: Despite previous in- tensive surveys for terrestrial snails in coastal New Britain, much of the diversity of the island’s interior mountains may remain to be described. More explora- tion is clearly needed in the Nakanai Mountains, where uplifted karst terrain has promoted the development of a unique and diverse snail fauna. Our cursory surveys, which did not penetrate deeply into, or reach high el- evations in the Nakanai Mountains, uncovered several previously unreported species, including the three Pary- phantopsis species described here. Most species of Pary- phantopsis appear to be restricted to single mountain ranges or islands. The absence of these three species from relatively intensively surveyed coastal areas suggest they are endemic to the Nakanai Mountains. It is likely that other endemic species will be found in the W hite- man, Baining, and Willaumez ranges of New Britain. The Charopidae were until recently, considered a minor com- ponent of the terrestrial mollusk fauna of New Guinea (Solem, 1983). Our surveys (Slapcinsky, 2005; 2006) sug- gest charopid species radiations on New Guinea and sur- rounding islands rival the spectacular radiations exhib- ited by this family in the oceanic Pacific (Solem, 1983). Based on shell and genital morphology, the relation- ships among the three Paryphantopsis species on New Britain are unclear. Paryphantopsis corolla and P. nu- Page 190 THE NAUTILUS, Vol. 121, No. 4 cella share several unusual traits that may indicate a com- mon origin. In both species, the marginal ectocones are divided into numerous irregular cusps, the base of the epiphallus is robust, and an apical diverticulum is present on the epiphallus. Paryphantopsis fragilicosta, on the other hand, lacks these characters but has a large, glo- bose shell that can be difficult to differentiate from that of P. nucella. However, all three New Britain species share unique rectangular bases to their marginal teeth, a possible syne spomorphy. If so, characters that have been historically used to determine monophyly, i.e. genital and shell morphology, appear to be rapidly evolving and vari- able in this group. Additional morphologic and genetic characters are necessary before a clear picture of the relationships between Paryphantopsis species within New Britain and the adjacent New Guinea mainland can be resolved. ACKNOWLEDGMENTS We thank the landowners of Marmar and Pomio for per- mission to work on their land and for field assistance: Anton Buntapeko, Fred Kraus, Esrum Lonpulpagetuna and Damian Matalo for additional field assistance; Gai- kovina Kula and Maureen Ewai of Conservation Inter- national for providing logistical support and advice; Florence Paisparea, Environmental Officer for East New Britain Province for facilitating our visit to Marmar; Papua New Guinea National Museum and Art Gallery for providing in-country collaborative assistance; Papua New Guinea Department of Environment and Conser- vation, Papua New Guinea National Research Institute, and East New Britain Provincial Government for per- mission to work in East New Britain Province; Karen Kelley, Electron Microscopy Core Laboratory, Univer- sity of Florida for imaging radulae; G. Paulay for lending photographic equipment; and Jack Worsfold for sharing bibliographic information. Fieldwork for this ee Was supporte od by National Science Foundation grant DEB 0103794 and the Unive rsity of Florida Foundation, McGinty Endowment. LITERATURE CITED Audra, P., S. E. Lauritzen, and P. Rochette. 2001. L’hyperkarst des montagnes Nakanai. Modéle d’évolution dun réseau juvénile (gouffre Muruk) basé sur des datations U/Th et paléomagnétiques des sediments. 93-99 in Audra, P., P. DeConinck, and J.-P. Sounier, (eds.) Nakanai 197S— 1998: 20 ans d’exploration, Association Hémisphére Sud, Antibes, 224 p Bavay, A, 1908. Mollusques terrestres et fluviatiles. Nova Sar Zoology 5: 269-292, pl. 14. Beehler, M. (ed.) ) 1993. A biodiversity analysis for Papua New Ge Papua New Guinea conservation needs as- sessment, Vol. 2. Biodiversity Support Program, Washing- ton, DC & Department of Environment and Conserva- tion, Boroko, Papua New Guinea, 1-433 p. Coen, G. S, 1922. Descrizione di nuovo specie di molluschi del Museo Civico di Genova. Annali del Museo Civico di Storia Naturale, Genova 9(3): 359-363. Flannery, T. F. 1995. Mammals of New Guinea. Cornell Uni- versity Press, Ithaca, 568 pp. Fulton, H. C. 1902. Descriptions of new species of land Mol- lusca from New Guinea. Annals and Magazine of Natural History 7(9): 182-154. Jutting, W. S. S. v. B. 1964. Non-marine Mollusca of West New Guinea. Part i Pulmonata, I. Nova Guinea, Zoology 26: 1-74, pls. 1-2 McAlpine, 1 R. fe Fryne, D, E. 2001. Land use change and intensification in Papua New Guinea 1975-1996. Asia Pa- cific Viewpoint 42(2/3): 209-215. Orenstein, R. I. 1976. Birds Ple esyumi Area, Central New Britain. The Condor, Vol. 78(3): 370-374. Paijmans, kK. 1976 Vegetation. ms Pajminas K. (ed.) New Guinea Vegetation, Australia National University Press, paper 212 pp. 1934, Systematische und tiergeographische Unter- ce iiber die Landschneckenfauna des Bismarck- Archipels. I. Archiv fiir Naturgeschichte 3(3): 445-488. Rensch, I, 1937. Systematische und ey ee Unter- suchungen iiber die Landschneckenfauna a Bismarck- Archipe els. IT. Archiv fiir Naturgeschichte 6(3); 526-644. Slapcinsky, J. 2005. Six new species of ere (Gas- tropoda: Pulmonata: Charopidae) from the Papuan Pen- insula of New Guinea. The Nautilus 119: 27-42. Slapcinsky, ]. 2006. Paryphantopsis (Gastropoda: Pulmonata: Charopidae) from the Louisiade Archipelago of New Guinea. The Nautilus 120; 119-130. Solem, A. G, 1959. On the family position of some Palau, New Guinea, and Queensland Jand snails. Archiv fiir Mol- luskenkunde $8: 151-155 pls. 12, 13, 2 figs. Solem, A. G. 1970. The Enc eee land snail genera Pilsbry- charopa and Paryphantopsis (Mollusca: Pulmonata). The Veliger 12: 239-264. Solem, A. G. 1983. Endodontoid land snails from Pacific Is- lands (Mollusca: Pulmonata: Sigmurethra). Part H, Fami- lies Punctidae and Charopidae, Zoogeography. Field Museum of Natural History, Chicago, ix + 336 pp. Thiele, J. 1925. Mollusken vom Bismark-Archipel, von Neu Guinea und Nachbar-Inseln. Zoologische Jahrbiicher 55: 119-146. Woodhead, J. D.. S. M. Eggins, and R. W. Johnson. 1998. Magma genesis in the New Britain Island Arc: Further insights into melting and mass transfer processes. Journal of Petrology 39: 1641-1668. Rensch, THE NAUTILUS 121(4):191-200, 2007 Page 191 Mysella gregaria new species, a bivalve (Galeommatoidea: Montacutidae) commensal with an intertidal burrowing sea anemone from North Carolina, USA Lennie Rotvit Thomas Fox Jorgen Littzen Ase Jespersen! Department of Cell Biology and Comparative Zoology Institute of Biology University of Copenhagen edie rsitetsparken 15 Dk- 2100 Copenhagen 0, DENMARK North Georgia College and State University Dahlonega, GA 30957 USA ABSTRACT Mysella gregaria new species (Bivalvia: Galeommatoidea: Mon- tacutidae) is described from Wrightsville Beach, North Caro- lina, USA. Several individuals were collected from the body column of an intertidal burrowing unidentified actinian. The morphology of the shell and soft parts is described and com- pared with other species of the genus from the W. Atlantic Ocean and with other bivalves associated with solitary antho- zoans. Mysella gregaria is a ctenidial brooder and specimens are either males or females; no true hermaphrodites were found. Contrary to many less social commensal bivalves, repro- duction in M. gregaria does not involve sperm storage. We suggest that this may be correlated with the species’ gregari- ousness. Additional Keywords: Mollusca, reproduction, anatomy, sperm morphology, gregariousness. INTRODUCTION During intertidal collecting on a muddy sand flat at Wrightsville Beach, North Carolina, USA, one of us (TF) collected a number of galeommatoidean bivalves from S—10 cm long specimens of an unidentified burrowing actinian. The host species was rare, as not more than an estimated 5-S specimens were found during occasional visits to the locality in the period from October 1970 through July 1975. Unfortunately, none of them were preserved for later identification. Only two of the col- lected actinians had commensal bivalves attached. Many bivalves of the superfamily Galeommatoidea have a commensal life style as they live together with species of bottom- dwelling marine invertebrate hosts such as other bivalves, polychaete s, sipunculans, ech- ' Corresponding author, ajespersen@bi.ku.dk iurans, crustaceans, and echinoderms. Commensal asso- ciations between bivalves and solitary anthozoans are re- stricted to three known cases (Yamamoto and Habe, 1961; Ponder, 1971; Oliver, 1993) and it was therefore judged to be of interest to examine the present associa- tion more closely. The study has disclosed that the bi- valves represent a new species of Mysella Angas, 1877, a genus that comprises both commensal and free- living mivalwes es and species that are commensal on certain con- ditions, free-living on other conditions (Ockelmann and Muus, 1978). MATERIALS AND METHODS Thirty-two bivalves were retrieved from the skin of a host specimen collected on 9 July 1975. The site of collection was a sandy mud tidal flat in Banks Channel, Wrightsville Beach, North Carolina, and the approximate coordinates are longitude 77.8° W and latitude 34.2° N. The anemo- nes were dug from the lower intertidal zone during a spring low tide. The number associated with the second host and the date of collection was not noted. The be- havior of the bivalves after removal from the host was not studied, but it was observed that they detached easily. Seventeen of the bivalves were preserved in Heiden- hain‘s Susa for several hours. This procedure decalcifies the shells and the sizes given based on measurements of the mantle are therefore approximate. Three other bi- valves were fixed in 70% ethanol and used for the de- scription of the shell and for type material. Six specimens were embedded in Araldite and cut into 2-jzm thick se- rial sections that were stained with toludine blue. Seven other specimens were embedded in Paraplast, sectioned, and the 8-jm thick serial sections stained with hema- toxylin and eosin (H+E). Ultrathin sections were per- formed on the testis of one of the males. These sections Page 192 THE NAUTILUS, Vol. 121, No. 4 were contrasted with uranyl acetate and lead citrate and examined in a JEOL 1LOOSX electron microscope. Scan- ning electron micrographs were made using a JEOL JSM-6335F SEM. Photos of the type specimens of My- sella casta (Verrill and Bush, 1897) (USNM 77632) and M. barbadensis Dall, 1899 (USNM 95703) were used for comparison with M. gregaria. Shell length (SL) and height (SH) are given to the nearest 0.1 mm. SYSTEMATICS Family Montacutidae Clark, 1855 Genus Mysella Angas, 1877 Mysella gregaria new species (Figures 1-20) (Figures 1-10): Observations The SL of twenty measured Description: SHELL were made on three shells. specimens varied from 3.0 to 6.0 mm. The measure- ments SLxSH in the type material are 4.5x3.6 mm, 5.1x3.8 mm, and 5.5x4.3 mm. The outline is almost per- fectly oval, slightly higher in the anterior part, all margins being evenly rounded. The valves are relatively flattened, very thin, semitransparent, and with a light- brown. to grey-brown periostracum, which is darkest in the dorsal part. There are no coatings of ferruginous deposits. The surface is smooth, with very fine commarginal lines and even finer radial striae. No growth checks were ever visible. The interior surfaces of the valves are polished. The umbos are not very prominent and placed slightly toward the posterior region. In the right valve there are two diverging teeth; the anterior tooth is elongate- subtriangular and more prominent than the narrower, posterior one. They are separated by a stout ligament placed immediately below umbo in a triangul: uv senliee The left valve is edentulous but has a produced dorsal margin that fits into the teeth of the right valve. The anterior adductor scar is subtriangular, the posterior more oval, both fused with the respective foot retractor scars. The pallial line is relatively broad and lacks a pallial sinus. MANTLE (Figure 11): The mantle folds are fused far behind and for a short distance to separate the mantle opening into an inhalant-pedal aperture and an exhalant aperture. While the first forms a long slit ma the four- fifths of the length of the ventral side, the exhalant ap- erture is very small and located far poste pai Since live animals were not observed, we do not know whether, or to which extent, the mid mantle fold may cover the out- side of the shell or whether there are any siphons. The mantle edges bear minute papillae that are most distinct Figures 1-4. Vysella gregaria. Cleaned shell of holotype l. rom inside. 4. Left shell seen from outside. Shell length Right shell seen from inside. 2. Right shell seen from outside. 3. Left 5.1. mm Rotvit et al., 2007 Page 193 Figures 5, 6. Mysella gregaria. Paratype, outside view of left (5) and right sides (6). Shell length 4.5 mm. in the dorso-anterior sector. A typical, ciliated rejection outline, whereas the smaller posterior adductor is more fold is located just anterior to the end of the inhalant- oval. A small pedal protractor muscle is located ventrally pedal opening. and clearly outside the anterior adductor muscle. The MUsCULATURE (Figures 11, 13): The anterior adductor two pedal retractor muscles are equally large and of muscle is the larger of the two and is subtriangular in moderate size. They terminate in the base of the foot, 9 Figures 7-10. Mysella gregaria. Holotype, SL.5.1 mm. Scanning electron micrographs of right hinge seen in direct view (7) and slightly tilted from below (8) and of left hinge seen in direct view (9) and slightly tilted from below (10). Scale bar represent 200 ym Page 194 THE NAUTILUS, Vol. 121, No. 4 Figures 11,12. Mysella gregaria. 11. Anatomy of male, right valve, mantle and gill removed, heart and pericardium omitted. 12. Light microscopic representation of sperm cell. Abbreviations: aa, anterior adductor muscle; ae, acrosome; ar, anterior pedal retractor muscle; bg, byssus gland; eg, cerebral ganglion; dg, digestive gland; f, flagellum; fg, foot groove; fu, fusion of left and right demibranch behind visceral mass; ga, gill axis: in, intestine; k, kidney; Ip, labial palps: mp, middlepiece; n, nucleus; pa, posterior adductor muscle: pg, pedal ganglion and statocysts; pp, protractor pedis muscle; pr, posterior pedal retractor muscle; so, sexual opening: ss, style sac; st, stomach; te, testis; vg, visceral ganglion. Arrowheads indicate water entering along the inhalant-pedal opening and exiting through the exhalant opening. while only a few fibers spread further into the foot. There are almost no muscle fibers serving the byssus gland. FOOT AND Byssus GLAND (Figures 11, 14-16): In pre- served specimens, the foot extends forward and a little beyond the shell margin. It is bluntly rounded in front. The ventral side is distinctly ciliated and has a furrow that extends from in front of the heel to tip of the foot. In some specimens a blackish pigment spot can be seen at the very tip of the foot. The very small byssus cavity appears Y-shaped in transversal sections as it consist of a right and left slit which both discharge into the median furrow by way of a single duct. The lining epithelium is composed of a mixture of ciliated and of mMUCOUS-pro- ducing goblet cells. A group of more distant glandular cells terminate between the cells of the byssus cavity epithelium through long, slender ducts, Although such cells are normally involved in the formation of byssal threads, none were ever seen either in sections or on vhole animals. The nature of the glandular epithelium indicates that slime threads, rather than normal byssal threads are produced by the byssus gland. GILLs (Figures 11, 17): The gill axis, which runs from near the umbo, has an oblique course backward. Each gill is triangular. An outer demibranch is entirely absent and only the inner one is present. The ascending lamel- lae of the left and right demibranch are fused behind the foot, more anteriorly to the lateral sides of the visceral mass. Both gills are fused to the mantle edges immedi- ately in front of the exhalant opening. A food groove is only present along the ventral edges of each gill. Inter- lamellar junctions are present, but they are very few, whereas interfilamentary bridges are numerous and oc- cur regularly, Right and left hypobranchial glands of nor- mal size occur well inside the exhalant aperture. A pair of relatively large labial palps lies on either side of the mouth, Their opposing surfaces are provided with 9-10 ciliated ridges. ALIMENTARY CANAL (Figure 11): The esophagus is a Rotvit et al., 2007 Page 195 Figures 13-16. Mysella gregaria. 13. Sagittal section of a male (SL 3.0 mm) showing the course of the protractor pedis muscle (pp). 14. Transverse section of a male (SL 4.3 mm) through the foot and the opening of the byssus cavity (be). 15. Byssus gland cells (bg) opening into byssus cavity 16. Transverse section through byssus cavity of the male shown in Fig. 14. Abbreviations: aa, anterior adductor muscle; ar, anterior pedal retractor muscle; be, byssus cavity; bd, byssus duct; bg, byssus glandular cells; eg, cerebral ganglion: ep, epithelium of byssus cavity: fg, foot groove: me, mucous cells. 2-4m thick Araldite sections stained with toluidine blue (13, 14 and 16) and $-ym thick paraplast sections stained with H+E (15). Scale bars represent 100 zm (13-15) and 30 zm (16). Page 196 THE NAUTILUS, Vol. 121, No. 4 2 eK a Deena rein OF POY hae a = oe Ota a pr Qo ag [Yee a es Sad Figures 17-20. Mysclla gregaria. Transversal section of male (SL 4.3 mm) (17) and of female (18, 19), 20. Testis with abortive oocyte (oo). Abbreviations: au, auricles; id, inner demibranch; in, intestine; k, kidney; ne, nerve; np, nephridiopore; ov, posterior wall of ovary; pe, pericardial cavity; pr, posterior pedal retractor muscle; rp, renopericardial canal; so, sexual opening; te, testis. 2-zm thick Araldite sections stained with toluidine blue. Scale bars represent 200 wim (17), LOOpm (18, 19) and 50 zm (20). Rotvit et al., 2007 Page 197 short curved tube. The stomach is relatively capacious and heavily cuticularized. The style sac forms a wide posterior continuation from the stomach, is elongated conical, nearly as long as the stomach, and directe d pos- tero- ventrally. It is placed within the right side of the visceral mass. The intestine leaves from the underside of the stomach near its connection to the stvle sac, runs alongside the style sac and forms a loop around its tip, then sie dorsally between left and right parts of the gonad to loop backwards to the rectum The bound: wy between the ciliated intestine and the unciliated rectum is marked by a sharp transition zone dorsal to the poste- rior adductor. The digestive gland is mainly located ventral to the stomach and around the posterior part of the esophagus. In addition to a large ventral communication with the stomach, there are also smaller openings into it. REPRODUCTIVE SysTEM (Figures 11, 17, 1S, 20): Eight sectioned bivalves (SL = 3.0-5.6 mm) were males, wile three other sectioned bivalves (SL = 4.1-6.0 mm) were females. No truly hermaphroditic specimens were found, but a few mature oocytes were present in the testis of one of the males (Figure 20). All males were sexually mature. The very large testis occupies the posterior half of the visceral mass being replaced more anteriorly by the style sac, stomach, ond digestive gland. The ge meral she ape as seen in decalcified specimens is relative ly constant. A large undivided pos- terior portion gives rise to right and left halves that sur- round the intestine and extend ventrally to send a few short branches forward. Two other forwardly directed finger-shaped branches embrace the stomach. The paired aaa issue from the most posterior, undi- vided part of the testis. They are short but have promi- nent funnel-shaped and heav ‘ly ciliated openings into the suprabranchial chamber. Spermatogenesis proceeds ev- erywhere along the walls of the different portions of the testis. There is no special chamber for storing the mature sperm which, in a non-orientated manner accumulate in huge numbers in the central parts of the different por- tions of the testis. Many spherical to ovoid cells, 5-7 ym in diameter, lie scattered among the mature sperm. These cells have a centrally place a nucleus plus one or two inclusions. We were unable to discover how these cells arise and if they are in the any way associated with the spermatogenesis. Because of the insufficient fixation the TEM micro- graphs were of low quality. They nevertheless show enough details to illustrate the general ultrastructure of the flagellate sperm cells (Figure 11, B). The acrosome is 1.4-1.5 pm long and basally (near the nucleus) 0.6 xm broad. It is divided into a terminal subspherical body (0.4x0.6 2m) and a slightly tapering acrosomal vesicle, which is basally deeply invaginated to accommodate a filamentous subacrosomal material. The nucleus is 0.7— 0.S xm across and 1.2—1.3 zm in length. The 1.4-1.5 wm long middle piece was extremely ill- -preserved without any identifiable mitochondria but seems from a broad base near the nucleus to taper backwards. The female sexual openings (Figure 18) are identical in size and structure to those of the male. Two females were obviously spent, but one of them had retained a few abortive oocytes (diameter ca. 60 jzm) within the ovary and a few embryos in the suprabranchial chamber. Ex- cept for this, none of six decalcified but unsexed speci- mens examined in transparent light (SL = 4.0-5.6 mm) were brooding ova or larvae. No structures for storing foreign sperm were found in any of the sectioned ts valves. EXCRETORY SYSTEM (Figures 11, 18, 19): Left and right halves of the kidney communicate in the median plane for a short distance. Each one is composed of several smaller and larger sacs with glandular walls. Antero- laterally the kidney opens directly to the suprabrancial chamber through two ciliated nephroducts, which are unchez wracteristically short and have porous nephrid- eae placed not ‘far from the genital opening. Rather far behind within the kidney, two long straight and heavily ciliated renopericardial canals run Ponand to open into the forward and ventral part of the pericar- dium. Host ReLations: All 32 individuals were attached onto the body column of a single sea anemone, whereas the number attached to a second host was not noted. None were found on any of the other host specimens from the same site, which suggests that the species is gregarious. It was observed that the attachment to the Rost was very loose as several of the bivalves had fallen off during the collection. This corresponds with the con- clusion that no true byssus threads are produced and that attachment rather takes place by means of slime threads. Holotype: BIV-445 (Zoological Museum, University of Copenhagen), a cleaned shell, SL = 5.1 mm. Paratypes: BIV-446, a shelled animal, SL = 4.5 mm; USNM _ 1107828 (National Museum of Natural History, Washington, DC), a shelled animal, SL = 5.5 mm; all from type locality, 9 July 1975. y: Banks Channel, Wrightsville Beach, North Carolina, USA. (ea. 77.8° W, 34.2° N), lower in- tertidal zone during spring low tide, sandy mud substra- tum. 9 July 1975. Thirty-two specimens were attached to a single host. Type Locality: Etymology: the species name is derived from the Latin grex, flock, and refers to the gregarious life style of the new species. DISCUSSION Identification: The details of the shell and hinge al- locate the species into the genus Mysella (family Mon- tacutidae) and indicate that it is close to the North At- lantic M. bidentata (Montagu, 1803). In M. gregaria the protractor pedis muscle is inserted immediately ventral to the anterior adductor as in M. bidentata (Montagu, 1803) and in Montacutona compacta (Gould, 1S61), in contrast to the condition in another group of mon- Page 195 THE NAUTILUS, Vol. 121, No. 4 tacutids, in which it splits the adductor in dorsal and ventral portions (Jespersen et al., in press). Mysella has often been combined with Rochefortia Velain, 1877, but the two differ with respect to the dimensions of the teeth of the right valve (Coan et al., 2000; Holmes et al., 2006). In species of Mysella the right valve has a stout trans- verse tooth anterior to the resilifer and a posterior tooth is either small or absent. In Rochefortia the right valve has two subequal diverging teeth, which smenuld place M. gregaria in Roc hefortia. The distinction ignores the av ailable soft anatomical characters which, especially in species with a Rochefortia type of hinge, vary consider- ably: A small outer demibranch is present in some (M. tumida (Carpenter, 1864), M. verrilli (Dall, 1899), and M. sovaliki MeGinitie 1959), absent in other (M. biden- tata and M. gregaria). The esophagus is a simple tube in all the species except for M. verrilli, in which it forms a suctorial proboscis. Foreign sperm are either not stored (M. gregaria and M. verrilli) ), attached directly to the gill surface (M. tumida), or stored in an unpaired pouch- shaped seminal receptacle within the visceral mass (M. bidentata) or in paired receptacles in the outer demi- branch (M. sovaliki). A new definition of the genus My- sella will probably be called for as additional data on the anatomy of a wider range of species will bring about a complete rearrangement of the genus and its included species. From the described species of Mysella from the East American waters, M. gregaria may be distinguished by the following chinacters: M. planulata (Stimpson, 1857) has a very prominent hinge and the umbo is placed more poste sriorly, It attaches to buoys and wharf pilings or oc- curs in muddy sand of the Zostera community (Abbott, 1974: Franz, 1973). In Mysella ovata (Jeffreys, 1881) the umbo is extremely protruding and - hoa far posteriorly. It occurs in de spths of 183 to 287 m. The shell of M. triquetra (Verrill and Bush, 1898) is ‘eualatee| and the poste ror part of the shell is distinctly rostrate, not ey venly rounded. Mysella verrilli, a deep water species, has the umbo placed far posteriorly and the esophagus is devel- oped as a suctorial proboscis (Allen, 2000). In M. stria- tula (Verrill and Bush, 1898), both teeth are delicate and very narrow, the anterior tooth is shorter, and the very small umbo is located more posteriorly than in M. gre- garia. The shell of the following two species were stuchier | from photos provided by the National Museum of Natu- ral History, Smithsonian Institution (USNM): The ante- rior part of a left valve of M. barbadensis Dall, 1899 (USNM 95703) is almost three times as long as the pos- terior part (in contrast to the claim that it is shorter as described by Dall, who obviously mistook a left valve for a right one). The left valve of the type of M. casta Verrill and Bush, 1898 from North Carolina (USNM 77632) is more elongate (SH:SL = 7:10) than the valves of M. gregaria (SH:SL = 7.8:10) and the anterior end relatively longer. Nevertheless, among the the shell of gregaria. ast American species, casta is the one most similar to M Comparison with North Atlantic/Arctic species of Mysella: Conchologically M. gregaria resembles M. bidentata (Montagu, 1803) but differs anatomically in that the latter has an unpaired seminal receptacle and dimorphic sperm (Jespersen and Liitzen, 2001). The shell of M. cuneata (Verrill and Bush, 1898) is distinctly asymmetrical, as the right valve shows a slight concavity with a consequent ait waren along the ventral margin (Gage, 1968). In M. tumidula ( (Jeffreys, 1866), the pos- terior shell margin is distinctly angular, not evenly rounded. Mysella moelleri (Morch, 187" 7) and M. sovaliki both differ from M. gregaria in the hinge structure and, more significantly, in having preserv ed a small outer demibranch (Petersen and Liitzen, in press). Mysella planata (Dall in Krause, 1885) has a thick shell, in which the left valve has the dorsal line modified into two teeth, and the right valve shows only one, anterior tooth (van Aartsen, 1996). Comparison with Other Bivalves Associated with Anthozoans: Commensalism between bivalves and anthozoans is rare. Nipponomontacuta actinariophila Yamamoto and Habe, 1961, is small Japanese bivalve that has been found attached immediately outside the ring of tentacles of Halcampella maxima Hertwig (Actiniaria: Halcampoididae). Details of the relationship are not known, except that three specimens sitting close together ren the ring of tentacles were ‘lhuatrated by "Hobe (1973). ). Although the outline of the shell and the position of the umbo in N. actinariophila are somewhat similar to the studied species, the hinges are clearly different, as teeth are only present in the left valve in N. actiniari- ophila, not, as in M. gregaria, on the right valve. Mon- tacutona ceriantha Ponder 1971, from "Cenanties sp. (Ceriantharia) in Moreton Bay, E Australia, is easily separated from M. gregaria in ‘that each valve has four cardinal teeth and a small outer demibranch is present. Furthermore, M. ceriantha is attached to the interior of the tube of the host, not to the body, and in a small numbers (Table 1) (Ponder, 1971). A third bivalve, Hal- campicola tenacis Oliver, 1993, from Rottnest Island off Perth, SW Australia has a hinge similar to Montacuta, the right valve with a strong anterior cardinal merging with a ee ginal lateral ridge and an obsolete posterior car- dinal. The left valve has anterior and posterior strongly projecting marginal extensions which fit into the right valve (Oliver, 1993). Besides, the ligament has a iitho- desma and there is a small outer demibranch. A single bivalve was found on each of six hosts (Halcampoides sp.., Actiniaria: Halcampoididae) among 20 collected. seems fairly obvious that all four anthozoan-associated bivalve species are not specifically interrelated | Reproduction: Eight of the sectioned bivalves (SL = 3.05.6 mm) were sexually mature males and three other (SL = 4.1-6.0) were females. The females were spent but one had retained a few embryos in the suprabranchial chamber which shows that this species, like other gale- ommiatoideans, is a ctenidial brooder. A few mature 0o0- cytes, left over from a previous ovulation, were present in Rotvit et al., 2007 Page 199 Table 1. Mean number of commensal bivalves per host specimen and reproductive specializations in montacutid bivalves. Abbre- viations: dw, dwart males; sp, spermatophores; sr, seminal receptables; ss, sperm sacs; tt, testis transplantation. Mean no. per Bivalve species host specimen Host species Repre rch ictive specializations References Montacuta percompressa ] Holothurioidea tt Fox et al., 2007 Peregrinamor ohshimai 1 Crustacea sr, dw Liitzen et al., 2001a Salpocola philippinensis ] Sipuncula tt? Liitzen et al., in press Halcampicola tenacis ] Anthozoa > pers. comm. (Graham Oliver) Litigiella pacifica 1-3 Sipuncula sr Liitzen and Kosuge, 2006 Montacutona ceriantha LS Anthozoa sr Ponder, 1971 Entovalva lessonothuriae <2 Holothurioidea sp Kato, 1998; Liitzen et al., 2005 Pythina arcuata ca. 2 Brachiopoda sr pers. comm. (JL) Anisodevonia ohshimai 1.8-2.7 Holoturioidea sp Kosuge, 2001; Liitzen et al., 2005 Nipponomysella subtruncata 2.5 Sipuncula sr, SS Liitzen et al., 2001b Tellimya ferruginosa <3 Echinoidea tt ob et al., 2007 Mioerycina coarctata 3.8 Sipuncula sr Gage, 1979 Mysella cuneata 5 Sipuncula sr Gage, 1968 one of the males, which could indicate that females may change sex and that the species shows her maphroditic tendencies. The species in all probability reproduces by outcrossing, since none of the sectioned bivalves showed truly simultaneous hermaphroditic characteristics. All galeommatoideans brood the ova in a ctenidial brooding chamber consisting of the inner and, if present, the outer demibranch. This is also where fertilization takes place. In a normally-filtering bivalve the gills and the currents they generate do not favor a casual jaiake of sperm suspended i in the water. The ciliary activity of the gills probably functions as a barrier for penetration of most sperm cells and the only other access to the brood- ing chamber is against the flow of filtered water expelled through the exhalant aperture. Many an have over- come these difficulties by bulk transfer of sperm in con- tainers of different nature to the female’s suprabranchial chamber (see O Foighil, 1985a, for a review). Exactly how they do this is not understood except in a single case (O Foighil, 1955b). Some species have minute dwarf males that are permanently and intimately associated with the much larger female and still other produce sperm of two types th at form spermatozeugmata. Some of that spermatozeugmata is probably capable of inde- pendent mobility. None of these methods of a precise transfer of sperm occur in M. gregaria. The testis is ex- ceptionally large in the species, mu we speculate that the resulting high production of sperm cells may compensate for the inevitable loss suffered during the transfer be- tween the opposite sexes of sperm cells that are simply broadcast into the water. Sperm transferred to a female (or hermaphrodite) are often stored for a considerable period either on the gills or in seminal receptacles of various types and location. e think it is likely that there may be some correlation between the presence or absence of storing devices and the chance of encountering bivalves of the opposite sex. The commensal galeommatoid bivalves are sequestered along with their host into a microhabitat that they prob- ably never leave. A few examples show that the number of montacutid bivalves present per host (or host burrow) is usually small (Table 1). Except for H. tenacis, which has not been anatomically studied, all these species have evolved various measures that allow long-time storage of sperm in the female (or hermaphrodite), which effec- tively enhance the chances that spawned ova can be fer- tilized even if no sexual partners are around. Conversely, in a highly social species, like M. gregaria, such mecha- nisms are evidently much less weeded: It would be in- teresting to study the relation between the sizes of the micro- populations i in other Montacutidae that neither ex- hibit sperm transfer nor sperm storage. Unfortunately, besides the present species, the known cases are limited to two species with unknown spatial distribution, Tel- limya tenella (Lovén, 1846) and Mysella moelleri (Fox et al. 2007; Petersen and Liitzen, in press). The present study has shown the need for further in- formation on the anatomy as a tool of a better under- standing of the taxonomy, which, to a much too large extent, has been based only on shell characters. More data are also wanted on the numerical relations between hosts and commensals and, above all, analyses on the sexual behavior of commensal bivalves are in very high demand. ACKNOWLEDGMENTS The study was supported by a grant to JL and AJ from the Danish National Science Found: ition (no. 51-00- 0278). We are grateful to Mr. Bjarne Bisballe, Zoological Museum, Cope snhagen, for operating the Scanning elec tron microscope aid to Mr. Gert Brovad, same institu- tion, for photographic assistance. We further thank Dr. Graham Oliver, National Museum of Wales, Cardiff, Uk for providing information about the occurrence of Hal- campicola tenacis on its host. LITERATURE CITED Abbott, R. T. American Seashells. 1974. 2nd edition. Van Nos- trand-Reinhold Co., New York, pp. 663 Page 200 THE NAUTILUS, Vol. 121, No. 4 Allen, J. A. 2000. An unusual suctorial montacutid bivalve from the deep Atlantic. Journal of the Marine Biological Asso- ciation of the United Kingdom S80; 827-834 Coan, E. V., P. V. Scott and F. R. Bernard. 2000. Bivalve Sea Shells of Western North America. 764 pp. Santa Barbara Museum of Natural History, Santa Barbara. Fox T. H., A. Jespersen and J. Liitzen. 2007, Sperm transter and reproductive biology in species of hermaphroditic bi- valves (Galeommatoidea: Montacutidae). Journal of Mor- ae 268: 936-952. Franz, D.R. 1973. The ecology and reproduction of a marine bivalve, Mysella planulata (Erycinacea). Biological Bulle- tin Woods Hole 144: 93-106. Gage, J. D. 1968. The mode of Mysella cuneata, a bivalve commensal with Phascolion strombi (Sipuncuidea). Cana- dian Journal of Zoology 46: 919-934. Gage, J. D. 1979. Mode of life and behaviour of Montecuta phascolionis, a bivalve commensal with the sipunculoidea Phascolion strombi. Journal of the Marine Biological As- sociation of the United Kingdom 59: 635-657. Habe, T. 1973. Halcampella maxima Hertwig, host of Nippono- montecuta actinariophila Yamamoto & Habe. Venus 31: 157. Holmes, A. M, J. Gallichan, and H. Wood. 2006, Coracuta obliquata n. gen. (Chaster, 1S97) (Bivalvia: Montacutidae) —first British record for 100 years. Journal of Conchology 39 (2): 151-158. Jespersen, Aand J. Liitzen. 2001. Ultrastructure of the seminal receptacle and the dimorphic sperm in the commensal bivalve Mysella bidentata (Veneroida; Galeommatoidea; Montacutidae). Acta Zoologica 82: 107-115. Jespersen, A J. Liitzen, and P. G. Oliver. In press. Morphology, biology and systematic position of Epilepton clarkiae (C lark. 1852) (Galeommatoidea: Montacutidae), a bivalve commensal with sipunculans. Journal of Conchology. Kato, M. 1995. Morphological and ecological adaptations in Montacutid bivalves endo- ectosymbiotic with holothuri- ans. Canadian Journal of Zoology 76: 1403-1410. Kosuge, T. 2001. Ecological notes on the populations of the galeommatid bivalve Anisodevonia ohshimai in Ishigaki Is- land, Southern Japan. Biological Magazine, Okinawa 39: 5-10. Liitzen, J., H. Sakamoto, A. Taguchi, and T, Takahashi. 2001a. Reproduction, dwarf males, sperm dimorphism, and life cycle in the commensal bivalve Peregrinamor ohshimai Shoji. Malacologia 43: 313-325. Liitzen, J.. T. Takahashi, and T. Yamaguchi. 2001b. Morphology and reproduction of Nipponomysella subtruncata (Yokoyama), a galeommatoidean bivalve commensal with the sipunculan Siphonosoma cumanense (Keferstein) in Japan. Journal of Zoology 245: 429-440. Liitzen, J. M. Kato, T. Kosuge, and D. O Foighil. 2005. Re- production involving spermatophores in four bivalve gen- era of the superfamily Galeommatoidea commensal with holothurians. Molluscan Research 25: 99-112. Liitzen, J. and T. Kosuge. 2006. Description of the bivalve Litigiella pacifica n. sp. (Heterodonta: Galeommatoidea: Lasaeidae), commensal with the Sipunculan Sipunculus nudus from the Ryukyu Islands, Japan. Venus 65: 193— 203. Liitzen, ]., T. Kosuge, and A. Jespersen. In press. Morphology of the bivalve Salpocola philippinensis (Habe and Kanazawa, 1981), new genus ( te eae Be La- saeidae), a commensal with the sipunculan Sipunculus nu- dus from Cebu Island, the Philippines. Venus. Ockelmann, K. and kK. Muus. 1978. The biology, ecology and behaviour . the bivalve Mysella bidentata (Montagu). Ophelia 17; 1-98. O Foighil, D. i Form function, and as of temporary dwarf males in Pse udopythina rugifera (Carpenter, 1864) (Bivalvia: Galeommatacea). The Ghicoe 27: 245-252. O Foighil, D. 1985b. Sperm transfer and storage in the brood- ing bivalve Mysella tumida. Biological Bulletin. Woods Hole 169: 602-614. Oliver, P. G. 1993. A new bivalve associated with a burrowing sea anemone. Halcampicola tenacis gen. sp. nov (Galeom- matoidea: Montacutidae) on Halc ampoide. ssp. (Anthozoa: Antipatharia: Halcampoididae). West Australian Museum, Perth, 1993: 255-265. Petersen, G, H. and J. Liitzen, In press. Morphology of Mysella moelleri (Mérch, 1877) and Mysella sovaliki MacGinitie, 1959 (Bivalvia: Galeommatoidea: Montacutidae), with notes on their biology. Steenstrupia, Copenhagen. Ponder, W. F. 1971. Montacutona ceriantha n.sp., a commen- sal leptonid bivalve living with Cerianthus. Journal de Conchyliologie, Paris 109: 15-25. Van Aartsen, J. J. 1996. Galeommatacea and Cymiacea. Part 2. Conchiglia 28: 27-53 Yamamoto T. and T. Habe. 1961. Nipponomontacuta actinari- ophila gen. et sp. nov., anew commensal bivalve of the sea anemone, Seto Marine Biological Laboratory 9: 265-266. THE NAUTILUS 121(4):201-209, 2007 Page 20] Corbula tarasconii, a new species of Corbulidae (Bivalvia) from offshore Brazil Eliane P. Arruda Osmar Domaneschi Departamento de Zoologia Instituto de Biociéncias Universidade de Sao Paulo Caixa Postal 11461 05422-970, Sado Paulo, BRAZIL arrudaep@yahoo.com.br domanesc@ib.usp.br Jonata de A. Francisco José Carlos N. de Barros Laboratorio de Malacologia Departamento de Pesca e Aqiticultura Universidade Federal Rural de Pernambuco Avenida Dom Manuel de Medeiros, S/N 52171-900, Recife, BRAZIL jonatal 98 1@yahoo.com.br mundovan4@yahoo.com.br ABSTRACT Corbula tarasconii, a new species of Corbulidae is described from material collected along the Brazilian coast. No living specimens are known and only shell characters were compared with the most similar Corbula species from the western Atlantic and eastern Pacific oceans. Short, ventrally curving rostrum and sculpture of low, rounded commarginal ribs he wing their bases about three times broader than the interspaces are the most important diagnostic characteristics of C. tarasconii. These characters distinguish the new species from other western At- lantic and eastern Pacific Corbulidae. Additional Keywords: Corbula tarasconii, Corbulidae, Bivalvia, taxonomy, Brazilian littoral, new species. INTRODUCTION During a research project carried out by the two senior authors on Corbula Bruguiére, 1797, occurring in Bra- zilian sea waters, specimens of an unknown species were found in samples made at four different localities be- tween southern Bahia state and northern Rio de Janeiro state. Working independently, the last two authors found the same unknown bivalve among the benthic fauna col- lected during field research on the continental shelf of northern Bahia state. Careful examination of all speci- mens gathered by the four authors, plus comparison with the nominal species hitherto published in the literature, led to us to conclude that the specimens represent a new species. Corbulids are readily recognized by their small size usually less than 20 mm in she ll length) and their in- equivalve condition, with the right valve larger, more convex, and overlapping the — one. All corbulids are shallow-burrowing suspension feeders inhabiting sandy, sandy-mud, or muddy substrata, usually at depths greater than 4 m Lamprell et al., 1998). Two living subfamilies of Corbulidae, Corbulinae Lamarck, 118, and Lentidiinae Vokes, 1945, encompass all extant (~S5) species; a few tropical representatives live in brackish rivers and streams (Coan et al., 2000). Len- tidiinae is represented by oan of the single genus Lentidium Cristofori and Jan, 1832: Corbulinae is by far more specious and includes 25 genus-level taxa (Coan et al., 2000), among which Corbula is the largest. Keen (1969) previously rubdaaded Corbula into 18 subgenera, several of which were rather poorly defined and in need of revision, whereas authors such as Warmke and Abbott (1961), Abbott (1974), Bernard et al. (1993), Coan et al. (2000), Mikkelsen and Bieler (2001) and Anderson and Roopnarine (2003) elevated some subgenera to generic status, a decision not shared by Coan (2002). In this latter paper, Coan (2002), considered elevating subgenera to genera premature, because the arrangement of these taxonomic categories is still fraught with inconsistencies and additional characters need to be better defined. The genus Corbula has long been a source of nomen- clatural confusion and many authors have been at- tempte -d to resolve it. It is beyond the s scope of this paper to discuss the systematics of the entire group once Coan (2002) has already presented a consensus based on the current rules of the Intemational Code of Zoological No- menclature (1999), According to Mikkelsen (2004), there are 13 species of Corbulidae in the western Atlantic, eight of which were previously cited by Rios (1994) as occurring on the Bra- zilian ap Corbula (Corbula) caribaea @ Orbigny, 1853, C. (C.) lyoni Pilsbry, 1897, C. ) patagonica dOrbigny, oe C. (C.) tryoni E. A. Smith, 1880, C. (Caryocorbula) sypmella Dall, hae: C. (Caryocorbula) dietziana C. B, Adams, 1852, C. (Juliacorbula) cubani- ana dOrbigny, 1853, and C. inate operculata Philippi, 1848. Revising Varicorbula from the western Atlantic, Mikkelsen and Bieler (2001) considered C. (V.) operculata cite od by Rios (1975; 1985; 1994) as a synony- mous with V. dis) yarilis (VOrb igny, 1842) or misidenti- fied specimens of V. philippit ( (E. A. Smith, 1885) Page 202 THE NAUTILUS, Vol. 121, No. 4 In this contribution, we describe a new species of Cor- bula from Brazilian waters, based on shell characters only, because no living specimens were obtained, and we compare this new species with its most closely related species from Atlantic and Pacific waters. MATERIALS AND METHODS Twenty whole shells and 56 disarticulated valves (28 right and 28 left) of the new species were collected on the Brazilian continental shelf, between northeastern Ba- hia state (11°58.7' S, 36°49.2' W), and northeastern Rio de Janeiro state (21°20'28” S, 40°16'09" W). Shell mor- phology was compared with the most closely related spe- cies known from the western Atlantic and eastern Pacific, borrowed from the Departamento de Zoologia da Uni- versidade de Sao Paulo, Brazil (one lot of C orbula aequi- valvis Philippi, 1836, and one of C. caribaea dOrbigny, 1853, both without catalog number), Museu de Zoologia a de Sao Paulo, Brazil ( C. bicarinata G. B. Sowerby, 1833, lot MZSP 67964), and Santa Barbara Museum of Natural History, USA (C. marmorata Hinds, 1843, lots SBMNH 83076, SBMNH 131640, and SB- MNH 141610). Shell characters and illustrations of C. ira Dall, 1908, provided by Coan (2002), were the basis for comparison with those in the new spe cies. The holotype and 14 paratypes were deposited in the malacological collection of the Museu de Zoologia da nixersidede de Sao Paulo (MZSP), 15 paratypes in the Museu Nacional do Rio de Janeiro (MNRJ), and 46 paratypes were deposited in the Museu Oceanografico Prof. Eliézer de Carvalho Rios (MORG). SYSTEMATICS Order Myoida Family Corbulidae Lamarck, 1818 Subfamily Corbulinae Lamarck, 1818 Genus Corbula Bruguiére, 1797 Subgenus Caryocorbula Gardner, 1926 Corbula tarasconii new species (Figures 1-15) Type Locality: Santo state, 20°45’ Off Guar: Municipality, Espirito S, 40°25 . Brazil, 60-65 m depth. Holotype: Museu de Zoologia, Universidade de Sao Paulo, MZSP 84452 (Figures 1-5, 8, 11). Measurements: 7 mm length, 5 mm height, 4 mm width Paratypes: MZSP 84453 to $4461, northeast of Bahia state, 11°58.7' S, 36°49.2' W., O1 Nov. 2000, 100 m (MZSP 84453: |] complete shell; MZSP 84454 to 84461: 6 right, 2 left valves); MZSP 84462 to 84464, southeast of Bahia state to - ar Best Municipality, Espirito Santo state, 15°33' S W to 20°45’ S, 40°25’ W (MZSP 84462-84464: 3 ee shells, 1 left valve); MZSP 86026, northeast of Rio de Janeiro state, 21°20'28" S, 40°16'09" W, Feb.—Mar. 1992, 139 m (1 complete shell); MNBRJ 11146-11157, northeast of Bahia state, 11°58.7 S, 36°49.2' W, OL ne 2000, 100 m (MNRJ 11146, MNBJ 11147: 2 complete shells; MNRJ 11148-11157: 5 right, 5 left valves); ); MNRJ 11040, southeast of Bahia state, 15°53'S2” S, 38°31'09" W, 30 Apr. 1996, 66 m (1 right, 1 left valve); ): MNRJ 11812, off Guarapari Munici- pality, Espirito Santo state, Oct. 1992, 60-70 m (1 com- plete shell); MORG 50792, off Boipeba Municipality, Bahia state, 13°35'18.33" S, 38°54'48.27" W, Feb. 2003, 41-53 m (4 complete shells); MORG 50789, off Camamu Municipality, Bahia state, 13°55'58.79" S, 38°05'28.13” W, 11 Dec. 2002, 52 m (7 complete shells, 16 right, 19 left valves). Diagnosis: Shell trigonal to trigonal-elongate, small (maximum length ~S mm), thin to moderately thick, in- flated, subequilateral, a a short rostrum, gently curv- ing ventrally. Left valve smaller than right. Sculpture sinailar in both shell valves, comprised of lowe regularly spaced rounded commarginal ribs crossed by minute, radially arranged pustules: base of commarginal ribs about three times broader than the anteroogll’ spaces. Description: Prodissoconch I and II markedly sub- orbicular. Prodissoconch I with a coarse and irregular surface texture under the SEM (length: 66.6 to 84.4 wm, n = 5); prodisocenct IT with subtle growth lines (length: 189.5 to 233.2 um, n = 5) and separated from the dis- soconch by a sharp transitional line and change in sculp- ture on the latter. Dissoconch small ( (length: min. = 2.4 mm, max. = 8.11 mm, mean = 5.92 + 1.36 SD [standard deviation]; height: min. = 1.7 mm, max. = 6.8 mm, mean = 4.48 + 1.06 SD; n = 47; measurement taken from right valve), trigonal to trigonal-elongate, moderately diisle in gerontic specimens, inflated, subequivalve, subequilat- eral with short rostrum. Free margin of the right valve completely overlapping the entire free margin of the left valve. Rostrum acutely rounded, gently curved ventrally. Posterior slope of each valve narrow, slightly concave, forming an acute angle (~20°) between posterior dorsal margin and the lew: rounded radial keel. Radial keel an inverted, gentle sig- moid line from umbo to the posterior limit of ventral margin; plane tangential to posterior slope forming a slightly obtuse angle with the plane tangential to cenrnl slope. Valve surface, excluding the posterior slope, regu- larly convex, except for a slight concavity in ee ventral area, just anterior to the keels. Umibos prosogy- rous, at about 36% of shell length from anterior end, aligned with cardinal tooth on right valve and cardinal socket on left valve. Anterior dorsal margin straight, ven- trally directed, continuous with evenly convex anterior margin, the latter situated below median longitudinal shell axis; posterior dorsal margin slightly convex, as long as, but less steep than the anterior dorsal margin; poste- rior margin long, obliquely truncated, and forming a short rostrum with posterior end of ventral margin; ven- tral margin evenly convex, except for a straight to slightly concave portion just anterior to the radial keel. Juvenile Arruda et al., 2007 Page 20 Ite mgnht va 3. Internal view of the left valve. 4. Internal view of the right valve. 5. Dorsal view to show the po keels (broad irrow) and escutcheon (narro irrow). Scale baz 2 mm 6. Paratype MNRJ 11154 external view of a transluce Page 204 THE NAUTILUS, Vol. 121, No. 4 Figures 7-S. a thin-shelled specimen (7) and a thick, gerontic specimen (8), showing differences in the development of the hinge plate at the resilial socket region (arrow). 7. Paratype MNRJ 11154. 8. holotype MZSP $4452. Scale bar = 0.5 mm. shells thin, whitish-translucent, turning moderately thick, and whitish-opaque as the specimens grow older; peri- ostracum partially preserved on posterior slope, espe- cially on left valve, missing on remaining shell area. External sculpture similar in both shell valves, com- prised of commarginal ribs crossed by minute pustules ~28.1 wm in basal diameter), the latter showing a ten- dency to align radially. ( onimarginal ribs regularly spaced, very low, rounded. with bases about three times broader than the intercostal spaces; commarginal ribs bec omimng moderate ‘ly elevated lamellae on poste rior slope of left valve only. Radial lines of pustules present all over shell surface, close »r to each other on poste nor slope: pustule lines visible through translucent shells. mediately \rea im- front of umbos sunken; lunule absent. Es- cutcheon lanceolate (~1/3 of shell leng h), delimited on right valve by a low, rounded elevation on the posterior slope, and on left valve by a slender radial rib; this slen- der radial rib formed by confluence and abrupt decrease in height of adjacent commarginal ribs, the last extending onto umbo almost parallel to free margin of escutcheon, Inner surface whitish porcelaneous, crowded with ran- domly scattered submicroscopic pustules (~16.5 jzm in basal diameter), distinguished under SEM only Hinge axis almost parallel to anterior dorsal margin. Right hing “sa with a cardinal tooth aligned with umbo, and a resilial socket sunken under umbonal lon: ¢ pe ae pik: ena stout with its apex curled clors: lly ISOSC( lc trian le shaped when viewed from its plat narrow deeply retracted at re Corbula tarasconii. Hinge of the right valve of Corbula tarasconii. Hinge of the left valve of Figures 9-11. thin- shelled specimens (9-10) and a thick, gerontic specimen (11), showing differences in the development of the hinge plate at the cardinal socket region (arrow). 9. Paratype MNRJ 11150; 10. Paratype MNBJ 11149; 11. Holotype MZSP $4452. Scale bar = 0.5 mm silial socket region in thin-shelled specimens, becoming expanded, thick, and more evident as specimens grow older. Left hinge plate with a deep, trigonal cardinal socket just posterior to umbo, and a thick, short chon- drophore projecting almost perpendicular to plane of hinge plate when viewed from its dorsal side; ee plate narrow, deeply retracted at cardinal socket region in thin-shelled specimens, becoming expanded, thick, and more evident as specimens grow older. Dorsal face of the chondrophore shallowly excavated and divided into an anterior and a posterior trigonal area by a radially placed, shallow, narrow groove; posterior margin of posterior trigonal area becoming thicker and projecting as a stout, rounded, tooth-like knob as specimens STOW older. Inner face of the right shell valve bearing a well- impressed commarginal groove for reception of entire free margin of the left valve. Anterior adductor muscle scar ovate, slightly to well-impressed; posterior adductor scar rounded in frontal view, on top of a slightly to well- elevated callosity. Anterior and posterior pe ‘dal muscle scars conspicuous and fused dorsally with corresponding adductor muscle scar. Pallial line narrow, glazed, far Arruda et al., 2007 Page 205 Figures 12-17. Scanning electron micrographs of Corbula tarasconii. 12-14. Paratype MZSP 84454. 12. External view of the left vi ei e showing regularly spe peed: low commarginal ribs (scale bar = 1 mm). 13. Frontal view of the posterior slope to show pustules radially arr inged and commarginal ribs turning into moderately elevate lamellae. Scale bar = 200 jum. 14. Detail of central slope (external view) showing the tendency of the pustules to arrange themselves radially. Scale bar = 200 jzm. 15-16. Paratype MZSP 84456. 15. Detail of the umbonal region and the hinge plate region of the right valve showing the low, rounded border of the escutcheon (arrow). Scale bar = 100 xm. 16. Internal view to show the presence of randomly seattered pustules. Scale bar = 50 jum. 17. Paratype MORG 50789, detail of the umbo to show the limit of both prodissoconch I ‘and II (arrows). Scale bar = 62 wm. from free border in both shell valves (farther in the right valve), especially in its anterior two thirds. Siphonal re- tractor muscle scar straight. Etymology: This species is named after Dr. José Car- los Tarasconi, a physician interested in collection and molluscan studies, who kindly donated the specimen from his collection, herein designated as the holotype. Observation: The holotype is the best preserved specimen among all complete ones; nevertheless, both shell valves hae. the postero-ventral portion of the ven- tral margin slightly broken. The brownish-red color viewed in the internal side of the holotype (Figures 3-4) was not observed among the paratypes. Remarks: The gay i in Corbula tarasconii of a small prodissoconch I (length: 66.6 to 84.4 jum), dis- Figure 18. Corbula tarasconii. Paratype MZSP $4462. Cam- tinctly separated from the larger prodissoconch I . : G 9339 5 era lucida drawing of the inner surface of the right valve show- (length: 189.5 to 233.2 pm), the latter devour of surface ing shell outlines, hinge, muscle scars and well impresse -d com- ornamentation, except for growth lines. 1 his suggests marginal groove for reception of the free margin of the oppo- that the species has planktotrophie development. accord- site valve (arrow). Scale bar = 1 mm. ing to the discussions in Jablonski and Lutz (1980) and THE NAUTILUS, Vol. 121, No. 4 Page 206 oor U1a}sva oylord UWIa}sea olor ULLayska onueyy UI9]SOA\ onUrpy UTOYSOA\ [izetg ‘onuR LY UTO}SOA\ O'T 99 AY uo porasya oIOUL “Sq dieys Aq pauyep asp Aq patttyop uu yuasqe quasqe sqtt oul squt divys Aq peuljep Use qUasqe qu lapueys Aq pouyep —AT Ajuo SuOQRAaa uowmal papunor jeuoquin “mor Aq suautoads oy uo peuryep sou quasaid —AY ut yudosaid ‘oqeprgsnd squt pareasya ‘divys Aq pauyep qu opus] Ss Aq peuyep me UONRAI]A pepano.t “axoy Aq peuyep —Ad yuosqe quosqe qUosqe aqyepnqsnd UWOHNLUSICL (Ulu) OZIs “XB N Woay wos soywyd erpeyy jeuoydig xodr aynor yy ‘squ ysuy ‘Buo.s squ a poept MOL “SuO.1S soords [eysoor9yUT ot} 0} yuapeatnba {APEX aseq pur xode oynor UIA “SqLt USTy A[eye.La pout 0} MOT soords [eJsoo.19}U OU 0} yuapeambe {PPE oseq TMV ‘sqLt papunos Ape.topout ‘MO[—A'T ‘soovds [eJsoo.19}Ut OU} ue} Lomoreu aseq UA ‘sqEt popunos 07 Iapus|s “MO[J—AY sooeds [eysoo.19} Ut Uy 0} yuapeamba (Pr aseq pur xed aynor UN “sque Ysty A[oye.lapOUuL OF AO] saoeds [eysoora}Ut ory wey TOPIAr aseq YIM “SqLt popunor “ANOT “OAT BA Ye = Al sre) ABA yu = AY ‘pouoquout you = Wu , pepuno.t cleys divys pepo. “AAC | yno}js ‘dieys popunor “A\OT SULAINO Al[e.quaa ‘poonpo.rd [Jo 0} Apetopout toys Hoy SIXe [Joys tottaysodo.1ajur oy (WU pousye ‘paoupord [pon 0} Ajoye.apout ploys SULAINO AyRAVUAA *}.0TS O}PAO-[RUOBLLL, aywapenbqus -9]PAO [PuosLy 0} ayapenbqus oWSUOTa -[PUOBLY Iypusurpaiqus aywsuola -[PUOSLY 0} RUOSLY aapeambaqn aageatmbaqns aageatnba aymb aapeammbaqns aapeamba oymb aaeambaqus DJVAOWADUL SD) DAL) DIDULIDIIG oO DIDGIADI “SD srapainbav “9 WUMOISVA [PULBAIRUTLLOD, aingdynog [poy WIMLSOY odeys ssouaapeamnbouy o (GOOG) UVOT) UO posed oIV DAr '—D JO SOYSLO~OVIBYE “SUBIdO OYLOR UWlo}sva puB OVUL|Y Uto}saM oY} WOLY sotods poryes Aposoyo S}t UL BSOY} TEM poardinoo NUOISVAD] “2 UL SOYSLOOVALY [POYS "LT Y[Quh Arruda et al., 2007 Page 207 Hain and Arnold (1992) on the relationships between prodissoconch morphology and modes of development. This new species encompasses all diagnostic charac- teristics presented by Keen (1969) and Coan et al. (2000) both for the family Corbulidae and genus em Its subequivalve, trigonal to trigonal- elongate shell with moderately coarse commarginal ribs, similar on both shell valves allow the inclusion of this species in the sub- genus Caryocorbula Gardner, 1926, as established by Anderson (1996) and Coan (2002). Beside these charac- teristics, C. tarasconii shares with Caryocorbula species a short chondrophore th that projects almost perpendicular to the plane of the hinge plate when viewed from its dorsal side. The allocation of C. tarasconii in Caryocorbula based in qualitative shell characters is an initial attempt to al- locate the new species to one of the named subgenera. As observed by Anderson and Roopnarine (2005), Caryocorbula “is relatively conservative in its morphol- ogy, making qualitative methods for alpha-level tax- onomy difficult.” Subgenera of Corbula have been poorly defined and fraught with inconsistencies, and a full-scale revision of the family is long overdue (Coan, 2002). Much more studies are needed to gather new taxonomic characters, which could better define the subgenus-level categories of Corbula and corroborate whether C, taras- conii is correctly allocated to Caryocorbula. Eighteen living species of Corbula are currently refer- able to the eastern Pacific (Coan, 2002) and thirteen to the western Atlantic (Mikkelsen, 2004). The western At- lantic species more closely related to C. tarasconii are C. aequivalvis and C. caribaea, and those of eastern Pacific water are C. bicarinata, C. marmorata, and C. ira. Corbula tarasconii is distinguished from C. aequiva- lvis by being approximately 50% smaller in length, con- spicuously inequivalve, with the posterior slope set off from the central slope by a low, rounded radial keel, sharp and stout in the latter species. The commarginal ribs in C. tarasconii are low, rounded, with the base about three times broader than the intercostal spaces and becoming lamellate on the posterior slope of the left shell valve, while in C. ae quivalvis they are low to mod- erately high, each with a quite acute apex and basal width equivalent to the intercostal spaces. The narrow, lan- ceolate escutcheon. better demarcated in the left shell valve of C. tarasconii, is another remarkable difference distinguishing this new species from C. aequivalvis, which has a wide, lanceolate escutcheon, wider in the right valve, and well-demarcated in both valves by el- evate ribs. The species can also be differentiated by the form and development of both the chondrophore and the right cardinal tooth. In C. tarasconii, the chondrophore has an inconspicuous to small tooth-like knob, and is shallowly excavated, with the dorsal face divided into two areas by a low, slender ridge; in C. aequivalvis, the tooth- like knob is lar ger and higher and the chondrophore is more aie from the free border of the hinge ai with the dorsal face divided into two areas by a high ridge, with the anterior area deeply excavated. Viewed from its convex face, the right cardinal tooth is right- triangle-shaped in C. aequiv alvis and isosceles-triangle- shaped in C. tarasconii. Corbula tarasconii greatly differs from C. caribaea by its short, ventrally curving rostrum that in C. caribaea is moderately to well produced and aligned with the an- tero- -posterior shell axis. Viewed from its inner surface, the posterior margin of the rostrum, in most individuals of the latter species, has a sinuous outline; the rostrum is frequently extended farther posterior by lateral, siphonal plates made of calcified periostracum. The rostrum of C. tarasconii neither has sinuous outline nor siphonal plates. The eastern Pacific C. bicarinata, compared to C, tarasconii, has an oval-subquadrate to trigonal outline, a shallow depression on the disc area farther anterior to the radial keel and aligned with the umbo-ventral axis, and the posterior slope set off from the central slope by a sharp, stout radial keel. The right cardinal tooth with a right-triangle shape and the wider, almost fan- shaped escutcheon, set off from the posterior slope by two stout, lateral ribs in C. bicarinata also differentiate it from C. tarasconii. Based on the figures and description given by Coan (2002), the eastern Pacific Corbula ira is similar to C. tarasconii in outline, configuration of the retractor sipho- nal muscle scar, umbos position, but greatly differs in its larger size and disc area sculptured vault strong, less nu- merous, rounded commarginal and fine radial ribs. Corbula marmorata, the third eastern Pacific species closely related to C. tarasconii, has a more elongate, trigonal-ovate shell sculptured with strong, high, and acute commarginal ribs anterior to the radial keel. Both species share a shallow depression just anterior to the posterior radial keel and a short, ventrally curved ros- trum, but the former species is also differentiated by the presence of a second shallow depression aligned with the umbo-ventral axis. ACKNOWLEDGMENTS The authors are greatly indebted to Paul Valentich-Scott, Santa Barbara Museum of Natural History, Santa Bar- bara, Luis R. L. de Simone, Museu de Zoologia da Uni- versidade de Sa0 Paulo (Brazil), Norma C. Salgado, Mu- seu Nacional, Rio de Janeiro (Brazil); and Paula Spo- torno de Oliveira, Museu Oceanografico Prof. Eliézer de Carvalho Rios (Brazil) for the loan of specimens of Cor- bula spp. Our thanks are due to Enio Matos and Eduardo Matos who provided assistance in the techniques of SEM, Departamento de Zoologia do Instituto de Bio- ciéncias da Universidade de Sao Paulo for scientific and infrastructural support, and FAPESP - Fundagao de Am- paro a Pesquisa do Estado de Sao Paulo that provided funding for this work. Special thanks to José H. Leal, Paula M. Mikkelsen and to the anonymous reviewer for their valuable comments on the manuscript. Page 208 THE NAUTILUS, Vol. 121, No. 4 LITERATURE CITED Abbott, R. T. 1974. American Seashells. 2"4 ed. Van Nostrand Company, 663 pp. Adams, C. B. 1852. Descriptions of new species of Corbula from Jamaica. Contributions to Conchology 12: 233-241. Anderson, L. 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Proceedings of the Zoological Society of London, 1833{1] (1-3): 34-38, Vokes, H. E. 1945. Supraspecitic groups of the pelecypod fam- ily Corbulidae. Bulletin of the American Museum of Natu- ral History, $6 (1): 1-32, pl. 1-4. Warmke, G. and R. T. Abbott. 1961. Caribbean seashells. A guide to the marine mollusks of Puerto Rico and other West Indian islands, Bermuda and the lower Florida Keys. Dover Publications, New York, 348 pp. THE NAUTILUS 121(4):210-213, 2007 Page 210 Two new gastropod species (Neogastropoda: Drilliidae, Turridae) from the western Atlantic Ocean Donn L. Tippett 10281 Gainsborough Rd. Potomac, MD 20854 USA ABSTRACT Two new deep-water species from the western Atlantic are proposed: Drillia (Clathrodrillia) blakensis and Hindsiclava rosensticlanus. Drillia blakensis is nearest Drillia (Clathrodril- lia) petuchi Tippett, 1995, and Hindsiclava rosensticlanus re- calls Hindsiclava polytorta (Dall, 1881). Animal anatomy, es- pecially foregut anatomy, is described for H. rosenstic anus. Additional Keywords: New species, Brazil INTRODUCTION The species proposed here are examples of the richness of the turrid fauna that continues to be discovered in the deep waters of the western Atlantic. Although collected in the 1960s, neither has been prey ously reporte sd. The type material of Drillia blakensis was part of the Bullis collection, secured as by-catch from the R/V OREGON. Hindsiclava rosenstielanus was dredged by the Univer- sity of Miami's R/V PILLsBury, but was only recently discovered during a review of the ane unsorted portion of the mollusk collection at the Miami's Rosenstiel School of Marine and pune Science. It is evident that further exploration and re- search will continue to reveal new material. MATERIALS AND METHODS Empty shells and shells with preserved animals were ex- amined. Preserved material was dissected. Radulae were mounted on microscopic slides and stained with Pronto- cil + CMCP 10, Type specimens were deposited at the National Museum of Natural History and other institu- tions. The classification used is that proposed by Taylor, Kantor, and Sysoev, 1993, which involved a rearrange- ment of the tre adition: I classification of the Turridae. Ab- breviations are: ANSP, Academy of Natural Sciences, Philadelphia Pe nnsylvania; MCZ, — um of Com- parative Zoology, Harvard Unive rsity, sachusetts: UMML School of Marine and Atmospheric Science, University of Miami, Florida; USNM, National Museum of Natural History, Smithsonian Institution, Washington, DC Cambridge, Mas- Invertebrate Museum, Rosenstiel Iniversity of SYSTEMATICS Drilliidae Olsson, 1964 Genus Drillia Gray, 1838 Type Species: Drillia umbilicata Gray, sequent designation, Gray, 1847. 1838, by sub- Subgenus Clathrodrillia Dall, 1918 Type Species: Murex gibbosus Born, 1778, by original designation. Drillia (Clathrodrillia) blakensis new species (Figures 1-3, $) ? Turridae sp.—Lamy and Pointier, 2001: 22, number 73, list, p. 26, photo. Description: Shell of medium-size (to approximately 45 mm), elongate, fusiform, with tall spire and large body whorl measuring about 0.5 shell length. Shell tapering gradually with moderate basal constriction to moderately elongate, open, slightly notched, slightly bent right ante- rior canal. Protosanch decollated, teleoconch w hols ten. Whorls well-rounded, shoulder sulcus on upper third concave, suture prominent. Sculpture of rounded, regu- larly spaced axial ribs with equal interspaces, e xtending from shoulder to followi ing whorl on spire and to base on body whorl. Ribs increase in number with shell growth, seven on early whorls, 12, narrower and closer spaced, on penultimate, ten or 11 on body whorl leading to small varix 0.25 whorl back of lip e dee, two or theese possibly abortive ribs following varix. Fine spiral threads overall, weaker on sulcus. Ape rture parallel-sided with apically directed, U-shi ape vd sinus poste poke bordered on body whorl by flat parietal tubercle. Lip sloping roundly for- ward below sinus, upper edge directed upward, narrow- ing sinus somewhat, producing spout-like appearance. Stromboid notch distinct. Color dirty-white overall, faint, pale brown peripheral band, blotch of same color on varix and spots preceding tops of axial ribs on later whorls. Operculum (Figure S) with roundly pointed anterior end and terminal nucleus. of chestnut color, ovate Type Material: Holotype, USNM 900034, 400450. mm, May 1965; three paratypes, USNM_ 1096708, data same Tippett, 2007 Figures 1-7. 44.5 x 13.6 mm, apertural, lateral, dorsal views. mm, off Cape San Antonio, Cuba. Scale bar = 25 mm. as for holotype, 45.7 x 16.6 mm, 44.1 x 14.7 mm, 42.5 » 16.0 mm (ex-José and Marcus Coltro collection., ex- author's collection); all dredged by R/V OREGON on type locality. Other Material Examined: elas Wolfe collection , 41.2 x OREGON on type locality. one specimen, Dr. Dou- 15.5 min, dredged by R/V Type Locality: Blake Plateau; precise location un- known, data presumably not retained. Distribution: Blake Plateau and possibly off Saba Is- land, Netherlands Antilles (Lamy and Pointier). Discussion: Drillia blakensis is most similar to Drillia Clathrodrillia) petuchi Tippett, 1995, from which it dif- fers in being narrower, having less robust ribs, finer spi- rals. a broader, non-tabulate, more sloping sulcus, and fainter color pattern. The author has not seen the shell figured in Lamy and Pointier, stated to be 48 mm in ength and from 150 m depth, however it appears from he illustration to be D. blakensis, differing in being slightly broader, having a slightly shorter anterior canal and stronger peripheral color banding, features within intraspecific variation limits. Etymology: Named after the Blake Plateau, the type ocality. Turridae H. Adams and A. Adams, 1853 (1838) Crassispirinae Morrison, 1966 Genus Hindsiclava Hertlein and Strong, 1955 Type Species: Clavatula militaris Hinds, 1543, by original designation. Hindsiclava rosenstielanus new species Figures 4-6, 9, 10 7. Pleurotoma (Drillia) polytorta Dall, 1881, holotype, Drilliids and turrids. 1-3. Drillia (Clathrodrillia) blakensis new species, holotype, USNM 900034, 44.8 «16.0 mm, apertural, lateral, dorsal views. 4-6. Hindsiclava rosensticlanus new species, holotype, USNM 1086746, off Riohacha, off Colombia, USNM 412171, 32.6 x 9.6 Crassispira polytorta (Dall, 1881): Okutani, 1983: 304, description and figures (apertural and lateral views plus radular teeth). ?Turridae sp.—Lamy and Pointier, 2001: 22, number 71, list and photo. rece iis 48 mm), elongate, fusiform, turreted, with tall spire, body whorl about 0.4 of shell length, whorls rounded below concave shoulder slope, gently constricted at base to moderately long, anterior canal open, notch lacking. Protoconch with 2.5 smooth whorls, tip central, first 1.5 whorls glossy, remainder dull-white, terminating in short portion of whorl with 4-5 axial riblets that quickly en- large to axial ribs in adult shell. Teleoconch whorls 9-10. Ape rture of parallel sides and with moderately deep, U-shaped posterior sinus on shoulder slope. Pé irietal tu- be rcle absent. Suture distinct, almost channeled, slightly Wavy. Subsutural cord of double threads. Axial ribs ro- bust (11-12 on body whorl, 11 on penultimate), extend- ing faintly across sulcus above and to next suture on Shell of medium-size (to approximately spire, disappearing on base; interspaces half again as wide. Uniform, regularly spaced spiral threads (5-6 on whorl periphery) cross ribs, producing modest, laterally elongate nodules, then continue forward to anterior ca- nal. Microsculpture of fine spiral threads, including sul- cus, irregular in strength and distribution, Notch and varix lacking. Color dirty-white overall, traces of dark periostracum., Gross Anatomy: Animal ee foot with upturned propodium operculum locate d posteriorly on foot Head bearing two tentacles, e me with eye dorsally on an co gee midway from base. Penis behind right ten tacle. reflected back under mantle Respiratory siphon on left a head, with moderate anal sinus on right. Gills and os bearing a fold, mantle edge extending across Page 212 THE NAUTILUS, Vol. 121, No. 4 Figures 8,9. Opercula, inner (left) and outer (right) views. 8. Drillia (Clathrodrillia) blakensis, 7.5 min length. 9. Hindsi- clava rosensticlanus, 11 mm length. Seale bar = 5 mm phradium visible through mantle. Rectum on right, Puckered rhynchostome between and slightly below ten- tacles. R hyne hostomal sphine ter present. R hynchocoel with strong linear folds internally. Rhynchodeal wall cir- cularly folded ef to retraction. Proboscis long, circularly folded along its length in preserved animal due to con- traction, linear folds posteriorly Large buecal mass and ivity posterio1 to rhynchodeum. No sphincter seen at beginning of esophagus. Large, highly coiled poison cland and bulb ‘lands present Radular ribbon ventral and posterior to body cavity. Right and left salivary Figure 10. Hindsiclava rosenstieclanus, radula, teeth are 300 um length. buccal mass. Gland ducts and radular ribbon enter at beginning of esophagus ventrallly and posteriorly to buc- cal mass. Incomplete radula an approximately 40 pairs of wishbone marginal teeth (Figure LO) measuring ap- proximately 300 jum. Operculum (Figure 9) medium am- ber, ovate with flat proximal side, rounded peripheral side, ends rather sharply rounded, terminal nucleus at anterior end, Type Material: Holotype, USNM 1086746, west of Riohacha, off Colombia, 11°32’ N 073°23' W, 549 R/V Pillsbury 781, 30 July 1968, 3 m, otter trawl, ex- UMML.30.10758; paratypes (ex-UMML,): one specimen, USNM 1107006; one specimen, MCZ 359155; one speci- men, ANSP 416320; seven specimens UMML 30.10758, four with animal preserved, three shells only. All from type locality. Other Material Examined: USNM 902064, three specimens, off Cartagena , Colombia, (ex-José and Mar- cus Coltro collection, ex-author’s collection); Pleuwrotoma (Drillia) polytorta Dall, 1SS1, USNM 412171, holotype. Type Locality: West of Riohacha, off Colombia. Discussion: Hindsiclava rosensticlanus is most similar to Hindsiclava polytorta (Dall, 1881) (Figure 7) reported from off Cuba, Hindsiclava rosensticlanus differs by its wider sulcus, doubled subsutural cord, fewer ribs (e.¢., nine on sixth spire whorl versus 12 on equivalent whorl of polytorta), presence of fine secondary spiral threads overall, and absence of parietal tube rcle. The specimen reported by Okutani, measuring 58 x 19 mm, from 328— 470 m off Surinam, appears to be this species despite the larger size and the radular teeth which are dissimilar at the location of their basal attachment (drawings appear stylized). The specimen reported by Lamy and Pointier (2001) from 450 in off Point Noire, Guade loupe, is 63 mm in length and, appears to be this species, again de- spite its larger size, and also despite the geogr: phic dlis- tance of Guadalupe from the type loc ality of rosenstiela- nus. The specimens in USNM 902064 are identical with the type mate rial exce pt that the axial ribs are slightly broader. Tippett, 2007 Page 213 Etymology: The species is named for the Rosenstiel School of Marine and Atmospheric Science, University of Miami, from which the eal was obtained. ACKNOWLEDGMENTS The author thanks the Department of Invertebrate Zo- ology, National Museum of Natural History, Smithsonian Institution for the opportunity of wor king with the mol- lusk collection and use of the equipment and facilities. Nancy Voss donated specimens of Hindisclava rosen- stielanus from the UMML. Yolanda Villacampa prepared the SEMs. Dr. Jerry Harasewych was constantly helpful in providing use of equipment, advice and support. The author thanks these people. LITERATURE CITED Dall, W. H. 1SS1. Reports on the results of dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico and in the Caribbean Sea, 1S77—79, by the U.S. coast steamer “Blake,” Lieutenant-commander C. D. Sigsbee, U.S.N., and Commander J. R. Bartlett, U.S.N., commanding. 15, Preliminary report on the mollusea. Bul- letin of the Museum of Comparative Zoology 15(2): 33- 144. Lamy, D, and J-P. Pointier. 2001. Les molluques profonds des Antilles Frangaises. Xenophora 95: 21-27. Okutani, T. 1983. Mollusks. In: Masatsune, T. and T. Okutani (eds.) Crustaceans and mollusks trawled off Suriname and French Guiana. Japan Marine Fishery Resource Research Center, pp. 187-354. Taylor, J. D., Yu. I. Kantor, and A. V. Sysoev. 1993. Foregut anatomy, feeding mechanisms, relationships and daseitt. cation of the C omoide sa (= Toxoglossa) mene eer Bul- letin of the Natural History Museum ( (Zoology) 59(2): 125- 170. Tippett, D. L. 1995. Taxonomic notes on the Western Atlantic Turridae (Gastropoda: Conoidea). The Nautilus 109: 127— 135. ; e + &, = ~ ; i THE@MNAUTILUS Volume 121 2007 AUTHOR INDEX ARRUIDA. Fi. IPs cote acdatmeactnin deed tratingunadseuankentiaenacehin 201 IIMAs Si Fe Bicwascseatecetancen arn dtwadagatdins aasutabiccsduh aan’ 99 BANDYOPADAYAY: Po Ks ccscscisveenesarsatetereaahe ovcuewesorataes 131 LOTZEN,, ewe inkses nenctaan tededdeanaa rontaesdsensatas scencatvanas 191 BARROS: [Guise ecsassneacsalsensseeisas tematssynaantesne 95, 99, 201 MARIOUIING, Py ote sitsctetaanssanitecesinie rangi eeselas srenteantes 159 BEU As Concpapetatonzatsutsaknbenuisannshddanaeeoasaaernns Scatenensn 90 IVECUINTAING Gz ne hcnkronnexsaaaniorranaaane noiathenwhocaunesnummaacete 139 BUYNEVICH Te Visca ccadisaastnccessactaliisessndinaiaradecercesnnes 37 INEKOILAS: [oacinedtinatactier dues aarinan sipinedssodmemmarenetnds 17 GOTEES alae He pita ty ieccheideta este eset tera decate cine ncatitadnuunsartatte eit eySars 17 OTIVERAS. Bist MMs siehose sa cweacacivsesesnss-aensarnsaitidoacdravns efetdle dean aR ncdlvetni a bu 13 GONCEPCION: (GPs asca Se aves Sanee sn eniosannanccneertesatoreestes 131 OWNBY), iePoicats hnensssegtwecetiae te cuou.ancs nettseta ricedihuawhde 13 POE VIRMES: Miva des ce aeteden cars hcmtete mia eonseaca taut agauamecanatatnaee 163 PEARCE,“ Ve fusauseiauessactenediirons aun teueteauewnteteeniaens 66 IDOMANESGHI, 'O% occ cinsccdexd ceteaenvasenh des thie cove seiabohenban 201 PENGHASZADEH: (Pie ss.4ccccetcnacoanactaantiecisagaidecavasasess 139 PRED Sy Mi Gi scccnosiianstana narakguaa adiscmeneainnonanuae Masten ceive 66 PETG, Ré Bi asnavoucsaviasendesstndnatcigestccans anadecaanenes 95, 99 NOs Wa ree cae Ae tea sce densi diese turecunamsniacdadanheatacd tena 191 PORTIS Ric Ws.53 vocaGanee gones-onigidtennalanssteiadatdacandeiaaens 105 IRANGISCO,, |). Asi. 2s.ctneneoametecesee vie doszetaie dnnmuatentesiae 201 ROTVIT:, Das iceland Co ncrua ne bed tenant oetonnd ade saakuuudeyaiteoeeoes 191 WREST D. Ye pcscceacsshteadhe2 teearianseideatais Moanenan tanner 43 ROY, Miviissesssicicisissusvorscctsvassaeristesiwnssasacsiesasieoeness 20 GAWEARDO, AG. Sizccdacacidcctaanavicassmeriaciesgwinnaagenaanenceeat 139 SANTOS SA DDS ss scsctaaesaincrancnaaaduinancad sent naewntaatincnd mi me 131 IARASEWYCH;, Mi (G..d.d esa ccneessassanctasanngadesave san 76, 90, 146 SAWE; Lisaas cvesacatnoussaantadgeesanida de odteah a concaamnadereaa avons 1 FIARDING, IM. oiiet oncedesenadsnsaan redisatecmacteatt aaageanssietons 146 SCHMELZ, G. W. 2... cc ccc cece cence eee e scenes eeenenesneeseenseees 105 EIERAEDE, Py Mi MT cenoncsesantnccoa vate unennmue oSeawead Wiecades 131 SLARGINSKY;. Joo. detawnssaacs dacs ihe senoneiasctee us tsaar cect nan ene 182 JESPERSEN, Ava ccsssecaccdsstaiiiadsensancarpncatensaateuhavas sabnnes 191 SQUIRES. Re Discs. diwasai shicwaricbidboiieas dest anercmdesieaances 1 INTEia 0: se anaes aabit weemacaseeoseenaah gaataeie heacecaminsaasenkaecans 76 STRONG 3 Eis: los dsasanainacasneacinhantradteniae tin manasncnsoecdet tinted 43 INURITA:: Kix foseucaSes toavawandeincd stuse cade wanendicaaceadieeaasawenes 66 PPE TAD Te Soon ii detehernt eae tee sciet sanewseae Reoeeee 210 TASTE: RS sole duciztsindemearsersittsts ie atssniernde seeadaednamatuenanenetelle 182 WY ATI =. ME secccc2 ccaateeete rade setae eine teins fied Gite awl aiaalatiarbiarae es 131 Pe MG Ss vec cat wn atest oaicuta ne apecauadsesaaaatnnnisaninanncoa te 104 WAEIAMSS WD TD ecincnspadsaawegeSedana socacd adn stwhcadsnah adeno 0 NEW TAXA PROPOSED IN VOLUME 121] GASTROPODA Bathrotomaria annejoffeae Harasewych and Kiel, 2007, new species (fossil, Pleurotomariidae).. 2... ee S4 Bathrotomaria bedetteae Harasewych and Kiel, 2007, new species (fossil, Pleurotomariidae) . 2... 82 Brachycythara beatriceae Mariottini, 2007, new species (Conidae) 0... ee 159 Cantallocostoma DeVries, 2007, new genus (fossil, Trochidae) .. 0... 169 Cantallocostoma panistostum DeVries, 2007, new species (fossil, Trochidae) .. 2... 0. ee 171 Chlorostoma quipua DeVries, 2007, new species (fossil, Trochidae). .. 0... 167 Cirsotrema chipolanum Schmelz and Portell, 2007, new species (Epitoniidae) .. 2... ee 110 Drillia blakensis Tippett, 2007, new species (Drilliidae) 2... 210 Epitonium conwaiae Schmelz and Portell, 2007, new species (Epitoniidae) .. 2... ee 122 Epitonium hoerleae Schmelz and Portell, 2007, new species (Epitoniidae). 2... eee 124 Epitonium incomitatum Schmelz and Portell, 2007, new species (Epitoniidae) Stata, Saas £0 ak Ue oes ee Bae es 120 Epitonium kalistos Schmelz and Portell, 2007, new species (Epitoniidae) . 2... 126 Epitonium regina Schmelz and Portell, 2007, new species (Epitoniidae). 0... 120 Epitonium vokesae Schmelz and Portell, 2007, new species (Epitoniidae) . 6... ee 126 Gerdiella alvesi Lima, Barros, and Petit, 2007, new species (Cancellariidae).. 2... . 100 Hindsiclava rosenstielanus Tippett, 2007, new species (Turridae) .. 2... 0 ee teats . 211 Intistoma DeVries, 2007, new genus (fossil, Trochidae) .. 2... ee eee 17] Intistoma pirqua DeVries, 2007, new species (fossil, Trochidae). 2... 0. ee ee a E72 Microcancilla jonasi Barros and Petit, 2007, new species (Cancellariidae)... 2... 02... boa) A emdvesr i BAL UE asitactated . 96 Paryphantopsis corolla Slapcinsky and Lasley, 2007, new species (Charopidae)..... 2.0... . Bees By & dine, Gos Bee are 183 Paryphantopsis fragilicosta Slapcinsky and Lasley, 2007, new species (Charopidae) . . . . . SG jun & Aone Gee pier _ 185 Paryphantopsis nucella Slapcinsky and Lasley, 2007, new species (Charopidae).. 6... 0. ee ee . 187 Obornella thompsonorum Harasewych and Kiel, 2007, new species (fossil, Pleurotomariidae)..... 6.2.0... Ais tits a ey MOO Opalia mica Schmelz and Portell, 2007, new species (Epitoniidae) ....... . eed GP URE ae aad pret teed Gate 2 3. da TTD Ee? IN A A WS Volume 121 2007 Opalia politesae Schmelz and Portell, 2007, new species (Epitoniidae).. 2... ee 115 Sassia melpangi Harasewych and Beu, 2007, new species (Ranellidae). 2... ee 90 Tegula masiasi DeVries, 2007, new species (fossil, Trochidae) .. 2.0... ee 175 Vertigo malleata Coles and Nekola, 2007, new species (Vertiginidae). 2... 19 BIVALVIA Corbula tarasconii Arruda, Domaneschi, Francisco, and Barros, new species (Corbulidae). .. 0... 0. ee ee 202 Mysella gregaria Rotvit, Liitzen, Jespersen, and Fox, new species (Montacutidae). 2. ee 192 Warren D. Allmon Rafael Araujo Jon Bryan Lyle Campbell Eugene V. Coan Robert H. Cowie Robert T. Dillon, Jr. Marien Faber Diarmaid O Foighil Matthias Glaubrecht Jochen Gerber M. G. Harasewych Kenneth Hayes REVIEWERS FOR VOLUME 121 Gregory Herbert Robert Hershler Steffen Kiel Frank Koehler Paula M. Mikkelsen Patricia Miloslavich Jeff Nekola Sven Nielsen Anton Oleynik P. Graham Oliver Marco Oliverio Guido Pastorino Kathryn E. Perez Sponsored in part by the State of Florida, Department of State, Division of Cultural Affairs, the Florida Arts Council and the National Endowment for the Arts v ENDOWMENT FOR THE ARTS Tra Richling Barry Roth John Slapcinsky Richard L. Squires Ellen E. Strong Jouni Taskinen Donn L. Tippett Paul Valentich-Scott David Véliz Geerat |. Vermeij Janice Voltzow Diego Zelaya iu a INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes articles on all aspects of the biology, Batenielony: and systematics of mollusks. Manuscripts describing original, unpublished research and review articles aati be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not require an abstract. Notices of interest to the mala- cological community will appear in a notices section. Manuscripts: Each original manuscript and accompa- nying illustrations Sena be submitted to the editor pref- alae y via e-mail or as hardcopy in triplicate. 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