Research 2006 Cainozoic , 4(1-2), pp. 71-96, February The Cantharus Group of Pisaniine Buccinid Gastropods: Review of the Oligocene to Recent Genera and Description of Some New Species of Gemophos and Hesperisternia ¹ Geerat+J. Vermeij ' Departmentof Geology, University ofCalifornia at Davis, One Shields Avenue, Davis, CA 95616, USA; e-mail: vermeij @geology,ucdavis. edu Received: 12 May 2004; revised version accepted 22 December 2004 The Cantharus of buccinid is in the Recent interval twelve of group pisaniine gastropods represented Oligocene to by genera, two which extinct. I review the and fossil record of these Anna 1826 are species composition, synonymy, characteristics, genera: Risso, (early Oligocene to Recent, eastern Atlantic); Cancellopollia Vermeij and Bouchet, 1998 (Recent, Indo-West Pacific); Cantharus Rdding, 1798 (Pliocene to Recent, Indo-West Pacific); Editharus Vermeij, 2001a (early Eocene to early Oligocene, Europe); Gemophos Olsson and Harbison, 1953 (late Miocene to Recent, tropical and subtropical America, one species in West Africa); Hes- peristernia Gardner, 1944 (late Oligocene to Recent, tropical and subtropical America); Pallia Gray in Sowerby, 1834 (early Mio- cene to Recent, Indo-West Pacific; one species in West Africa); Preangeria Martin, 1921 (early Miocene to Recent, Indo-West Pa- cific); Prodotia Dali, 1924 (?late Miocene to Recent, Indo-West Pacific); Pusio Gray in Griffith and Pidgeon, 1834 (?early and middle Miocene, late Miocene to Recent, eastern Pacific); Solenosteira Dali, 1890 (late Miocene to Recent, tropical America); and Zeapollia Finlay, 1927 (Oligocene to Pliocene, Australia and New Zealand). Besides many generic reassignments, I describe the basidentatus Pleistocene, Pliocene, following new species: Gemophos (early Florida); G. crispatus (late Florida); G. filistriatus (Recent, North Carolina to eastern Florida); Hesperisternia binodosa (late Pliocene, Atlantic Costa Rica); H. distans (early Mio- cene, Venezuela); and H. petuchi (late Miocene, Maryland). New morphological data are presented for Gemophos tridentatus (Tuomey and Holmes, 1857) from the late Pliocene ofthe southeastern United States. other shallow-water molluscan clades in its in Biogeographically, the Cantharus group conforms to many having highest diversity the Indo-West Pacific (about 25 species), followed by the eastern Pacific (19), western Atlantic (7), and eastern Atlantic (5). Within the western Atlantic, a distinction at the species level between the Gatunian Province (Greater Antilles to Brazil) and the Caloosa- hatchian Province (southeastern United States) has existed throughout the Neogene. At least three invasions from the Gatunian to the Caloosahatchian Province are documented, two in Gemophos and one in Solenosteira. In addition, Gemophos spread across the West and Pallia extended its from the Indian Ocean West Africa. intense Atlantic to Africa, range to Neogene extinction was most in the western Atlantic, where the end of the Pliocene witnessed the regional or global extinction of at least two lineages of Sole- nosteira and each and members of the Cantharus in the Plio- one of Gemophos Hesperisternia. Warm-temperate group disappeared Mediterra- cene and Pleistocene in California,Australia, New Zealand, and the northwestern Atlantic, and survive today only in the nean. Key words:: Buccinidae, Neogene, Gastropoda, taxonomy, biogeography. Introduction the genus-level taxonomy, morphology, biogeography, and evolution of the Cantharus group of pisaniine hue- The history of the marine tropical biota is a complex tale cinid gastropods, and I complement a species-level of evolution, invasion, and extinction against a backdrop summary for most of the genera with more detailed ac- tectonic of effec- of American and of climatic and change. One the more counts two genera, Gemophos Hes- tive tactics in telling this history is to interrogate the spe- peristemia. cies taking part in it. This, in turn, requires a detailed of the and rela- understanding taxonomy phylogenetic tionships of clades that have achieved a circumtropical Systematic Palaeontology distribution and for which a substantial fossil record is available. Abbreviations - The following abbreviations are used to This is contribution that end. In I examine referred in the paper a to it, denote repositories for material to text. -72- ANSP Academy of Natural Sciences, Philadelphia, phylogenetic relationships among these groups. PA, USA I twelve currently recognize genera in the Cantharus NMB Naturhistorisches Museum, Basel, Switzerland with These group post-Eocene representatives. are Anna, PRI Paleontological Research Institute, Ithaca, Cancellopollia, Cantharus, Editharus, Gemophos, Hes- NY, USA peristernia, Pallia, Preangeria, Prodotia, Pusio, Sole- RGM Rijksmuseum voor Geologic en Mineralogie nosteira, and Zeapollia. Preangeria, an early Miocene to (now Nationaal Museum His- Recent of four fossil voor Natuurlijke genus and one Recent species, di- toric), Leiden, The Netherlands verges widely from other members of the Cantharus TU Tulane New Orleans, LA, USA University, group, and was discussed by Vermeij (1998a, 2001b); I UCMP University of California Museum of Paleon- include it here for Prodotia only completeness. may be tology, Berkeley, CA, USA more closely related to the Pisania group. Although I UF University of Florida (now FloridaMuseum of shall characterize it here, its briefly taxonomy and scope Natural Gainesville, FL, USA research; I therefore History), require more confine myself to a USNM United States National Museum of Natural general comparison to other genera. History (Smithsonian Institution), Washing- ton, DC, USA Genus Anna Risso, 1826 Buccinidae 1815 - Anna Family Rafinesque, Type species massena Risso, 1826,by monotypy. Subfamily Pisaniinae Gray, 1857 Remarks - Based on my examination of most of the de- Remarks - As understood by the scribed the Vermeij (2001a), species, genus Anna is characterized by a Pisaniinae of shallow- comprises a group mainly tropical, fusiform, basally weakly constricted shell; weak to obso- water buccinid hard bot- lete predatory gastropods living on axial sculpture of 11 or more ribs on the last teleo- in toms, seagrass meadows, and on firm conch weak cords occasionally whorl; ten or more spiral or threads, sand. The shell is small to medium-sized for the often with threads family between and on the cords, on the last (maximum about 100 mm), ovate to fusiform, with teleoconch distinct length whorl; a parietal tooth at the adapical a short canal, crenulated outer rounded axial end of the inner siphonal lip, lip; a terminal, varix-like outer lip with ribs (when and a columella with two present), one or downwardly convex, crenate edge with the crenations weak basal folds. The rachidian of the radula square unpaired and with six or more denticles or short lirae on bears three typically with fusiform bases, the cen- its inner side; low fasciole often cusps siphonal present; um- tral generally being the largest and the lateral bilical chink absent. cusp cusps, No species I have seen exceeds a which sometimes smaller bifurcate, being (see Cemohor- of 27 mm. The from length genus ranges the early Oligo- 1971, 1975). The Pisaniinae extend back to sky, certainly cene to the Recent in the Mediterranean, western Europe, the late Cretaceous (Campanian), and possibly into the and West Africa. Cretaceous early (see Vermeij, 2001a). Well-known species include A. assimilis (Reeve, 1846), Using Cernohorsky’s (1971, 1975) scheme of classifica- Recent, Senegal; A. badensis (Hornes and Auinger, tion as a point of departure, I divide the Pisaniinae in- Badenian middle 1890) = Langhian Miocene, Vienna into three which formally groups, may or may not be Basin; A. bredai (Michelotti, 1847), Tortonian late Mio- clades These the Pisania (Vermeij, 2001a). are (1) cene, Italy; A. cancellarioides (Basterot, 1825), Aquita- group, comprising Eocene to Recent species with gener- nian and Burdigalian early Miocene, southern Europe; A. thin shells terminal ally having a varix, fine to obsolete consobrina (Cossmann and Lambert, 1884), early Oligo- adult reduced axial spiral sculpture, sculpture, and a high cene, France; A. dorbignyi (Payraudeau, 1826), late the also from the Eo- spire; (2) Engina group, ranging Miocene to Recent, Mediterraneanregion and Paratethys cene to the Recent, a narrow denticles with having aperture, (likely many synonyms named by Bellardi, 1873; on the inner side of the terminal and Homes and (adaxial) outer lip, a Auinger, 1890; see De Gregorio, 1885); and distinct axial ribs the and varix, on teleoconch; (3) A. minutulus (Baluk, 1995), Tortonian late Miocene, the Cantharus from the Cretaceous on- group, ranging Poland; A. massena Risso, 1826 (= Buccinum scac- ward, in which the inner side of the is chianum outer lip usually Philippi, 1844; see Dali, 1918; Arnaud, 1977), lirate rather than the is denticulate, aperture broadly Recent, Mediterranean; A. scabra (Locard, 1886), Re- the adult with with- ovate to ovate-elongate, outer lip or cent, Mediterranean; and A. unifilosa (Bellardi, 1873), terminal and and out a varix; spiral axial sculpture is Tortonian late Miocene, Italy. well on the teleoconch. Most fossil of usually developed species Anna were assigned by their de- In earlier I treated the an paper (Vermeij, 2001a), pre- scribes to Pallia. Anna differs from the Indo-West- members of the Cantharus Here, the Pacific Oligocene group. (IWP) genus Pallia by lacking a labral tooth at focus shifts to evolved