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Gastropoda: Turbinellidae) Ruthenica, 200 I, II (2): 81-136. ©Ruthenica, 2001 A revision of the Recent species of Exilia, formerly Benthovoluta (Gastropoda: Turbinellidae) I 2 3 Yuri I. KANTOR , Philippe BOUCHET , Anton OLEINIK 1 A.N. Severtzov Institute of Problems of Evolution of the Russian Academy of Sciences, Leninski prosp. 33, Moscow 117071, RUSSIA; 2 Museum national d'Histoire naturelle, 55, Rue BufJon, 75005 Paris, FRANCE; 3 Department of Geography & Geology Florida Atlantic University, 777 Glades Rd, Physical Sciences Building, PS 336, Boca Raton FL 33431-0991, USA ABSTRACT. The range of shell characters (overall established among some of these nominal taxa. shape, sculpture, columellar plaits, protoconchs) Schematically, Exilia Conrad, 1860, Palaeorhaphis exhibited by fossil and Recent species placed in Stewart, 1927, and Graphidula Stephenson, 1941 Exilia Conrad, 1860, Mitraefusus Bellardi, 1873, are currently used as valid genera for Late Creta­ Mesorhytis Meek, 1876, Surculina Dall, 1908, Phe­ ceous to Neogene fossils; and Surculina Dall, 1908 nacoptygma Dall, 1918, Palaeorhaphis Stewart, 1927, and Benthovoluta Kuroda et Habe, 1950 are cur­ Zexilia Finlay, 1926, Graphidula Stephenson, 1941, rently used as valid genera for Recent deep-water Benthovoluta Kuroda et Habe, 1950, and Chatha­ species from middle to low latitudes. Each of these midia Dell, 1956 and the anatomy of the Recent nominal taxa has had a complex history of family species precludes separation of more than one genus. allocation, which has not facilitated comparisons Consequently all of these nominal genera are sy­ on a broader scale. Exilia and Benthovoluta are the nonymised with Exilia, with a stratigraphical range genera best known in the fossil and Recent litera­ from Late Cretaceous to Recent. Anatomically, ture, respectively. Exilia is similar to other ptychatractine genera, With the exception of Benthovoluta c1aydoni but is characterized by a stomach with a long, Harasewych, 1987, which was taken in moderate narrow caecum, a penis with terminal fold sur­ quantity in the 1980s as a by-product of shrimp rounding the seminal papilla, and a radula with trawling off Western Australia, material of Bentho­ rachidian teeth with broad lateral flaps. Recent voluta and Surculina is rare in museum collections species of Exilia are restricted to deep water at and this, in turn, has not facilitated an analysis of middle to low latitudes in the Indian and Pacific the systematics of the group, which can be sum­ oceans. Exilia hilgendorfi (Martens, 1897) is treated marized as follows: as a species highly variable within its broad Indo­ Pacific distribution, with Benthovoluta gracilior Reh­ (I) At family level. At the time of its estab­ der, 1967, B. c1aydoni Harasewych, 1987, and B. lishment by Kuroda and Habe [1950], Benthovoluta prellei Bozzetti, 200 I considered local variants. was then classified in the family Volutidae. Based Three new species are described: Exilia graphidu­ on Habe's [1952] very schematic illustrations of loides sp. nov. (New Caledonia, 520 m), E. vagrans the radula, it was transferred to Turbinellidae by sp. nov. (West and SW Pacific, 865-1280 m), Kuroda [1965]. Harasewych [1987] described the and E. kiwi sp. nov. (New Zealand, 1386-1676 anatomy of B. c1aydoni and placed Benthovoluta in m). the subfamily Ptychatractinae of the Turbinellidae. Surculina was transferred from the Turridae to the Turbinellidae by Rehder r1967). Introduction (2) At genus level. Rehder [1967) discussed the relationships of Benthovoluta with Surculina and The species- and genus-level taxa discussed in kept the two genera as valid, and this has been the this paper have, at one time or another, been accepted genus-level taxonomy to this day. classified in half a dozen different neogastropod families, from Fasciolariidae to Volutidae, through (3) At species level. Thirteen Recent nominal Turridae and Turbinellidae, depending on the re­ species have, at one time or another, been attributed lative importance given to such shell characters as to Benthovoluta. Eight of these were transferred to columellar plaits and subsutural concave sulcus, or Cyomeslis Quinn, 1981 [Harasewych, 1987], now to anatomical characters when these became a synonym of Latiromitra Locard, 1897 [Bouchet known. No less than 10 nominal genera were and Waren, 1985), or to other genera [Bouchet established to accomodate the fossil and Recent and Kantor, 2000], and five species are currently species considered. Synonymies had already been recognized as valid. In addition, four species are 82 Yu. Kantor, Ph. Bouchet, A. Oleinik regarded as valid in Surculina [Rehder, 1967; Cer­ The fossil and Recent nohorsky, 1973]. nominal genera The present paper was prompted by the collec­ ting of new deep-water material attributable to the Genus Exilia Conrad, 1860 "Benthovoluta-Surculina" complex, which demon­ strates a more diverse range of shell shapes than Exilia Conrad, 1860: 291. earlier described and provide new anatomical ma­ Type species (by monotypy): Exilia pergracilis terial that permit a re-evaluation of genus-level Conrad, 1860, Midway Group, Eocene, Alabama, relationships. This has necessitated a review of USA (Fig. lA-D). several Recent and fossil nominal genera with Diagnosis: Shell narrowly fusiform, slender; aper­ slender fusiform shells and columellar folds. As a ture narrowly elongated; anterior canal long, stra­ result, we have found that Recent Benthovoluta is ight, and narrow; columella may be smooth or bear not generically separable from Late Cretaceous to one to four plaits of variable strength; surface with Neogene fossils and that the oldest generic name well developed axial and spiral sculpture; proto­ available for this group is Exilia. In this paper, we conch smooth, paucispiral or multispiral, depen­ discuss the generic synonymy and justify the use ding on the type of larval development. of Exilia and we describe the anatomy and revise Operculum from medium-sized with terminal the species-level systematics of the Recent taxa. A nucleus to very small with subcentral nucleus, or revision of the species-level systematics of the fossil absent. Proboscis vely short, with basal buccal mass; taxa was outside the scope of our work. ventral odontophore retractor passing through the nerve ring; single accessory salivary gland; large Abbreviations and text conventions: noe - nnterior and bulky gland of Leiblein; stomach with narrow oesophngus; nsg - nccessory snlivmy glnnd; cne - cnecul1l of stomnch; cl1le - cut mnntle edge; col.m - colul1lellnr and long caecum; penis with seminal papilla sur­ muscle; ct - ctenidium; dd - dend collected specimen; rounded by a nearly circular fold. Rachidian radular ddg - dUC;l of digestive glnnd; dg - digestive gland; teeth with broad lateral flaps. dgL - duct of glnnd of Leiblein; gL - gland of Leiblein; Remarks: Exilia is rather distinct from the other hg - hypobmchinl glnnd; Iv - live collected specimen; known Recent genera of Ptychatractinae in its moe - mid-oesophngus; ne - nephridium; ng - neph­ generally narrow fusiform shell with long and stra­ ridinl glnnd; nr - nerve ring; od - odontophore; op ight siphonal canal. Specimens of E. expeditionis - opcrculum; os - osphradium; p - penis; poe ­ may superficially resemble Met<geria alba (Jeffreys, posterior oesophagus; pI' - proboscis; PI'S - rhyncho­ 1873), the type species of Met:<geria Norman, 1879; daeum (= proboscis sheath); I' - rectum; s - siphon; the rachidian tooth of the radula of the latter, illus­ sd - salivary duct; sem.gr - open seminal groove; sem.p - seminal papilla; sg - salivary gland; st ­ trated by Bouchet & Waren [1985: fig. 393], differs stomach; t - head tentacle; tes - testis; vL - valve by having no lateral flaps. Species of Exilia also of Leiblein; vodr - ventral odontophore retractor; vpr ­ supeIiicially resemble species of Exilioidea Grant & ventral proboscis retractor. Gale, 1931, transfered from Buccinidae to Turbinel­ lidae by Bouchet & Waren [1988]. The anatomy of Repositories: AMNH - American Museum of atural Exilioidea is not known but, judging from its radula, History, New York; AMS - Australian Museum, Sydney; DMNH - Delaware Museum of Natural History, Green­ it also belongs to Ptychatractinae. The velY minute ville, USA; EAS PU - Department of Eanh and At­ size of the ribbon (less than 140 ~tm in length) and mosperic Sciences, Purdue University, FIOIida, USA; MNHN the narrowly arched, almost wishbone-shaped, central - Museum national d'Histoire naturelle, Paris; NM ­ tooth, with socketed central cusps interlocking on Natal Museum, Pietermaritzburg, South Africa; NM NZ successive rows, are unique to this genus. - Museum of New Zealand, Te Papa Tangarewa, Wel­ Anatomically Recent species of Exilia are rather lington; NZOI - New Zealand Oceanographic Institute, uniform, while they differ from other Ptychatrac­ Wellington (now NIWA); PRI - Paleontologicnl Resenrch tinae in having a large bulky gland of Leiblein, a Institution. Ithaca, USA; UCBMP - Museum of Pa­ radula with rachidian teeth with broad lateral flaps, leontology, University of California, Berkeley, USA; a stomach with narrow and well pronounced cae­ USGS - United States Geological Survey; USNM ­ National Museum of Nntural History, Smithsonian In­ cum, and a penis with seminal papilla surrounded stitution, Washington, DC, USA: UWBM - The Burke by a nearly circular fold. Museum, University of Washington, USA; WAM - Wes­ The family allocation of Exilia has fluctuated tern Australian Museum, Perth. considerably until quite recently.
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