Author's personal copy Pala¨ontol Z (2012) 86:451–456 DOI 10.1007/s12542-012-0145-1
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The oldest Odontostomidae (Mollusca: Gastropoda): Bahiensis priscus n. sp. (Paleocene, Uruguay) Fernanda Cabrera • Sergio Martı´nez
Received: 13 April 2012 / Accepted: 13 June 2012 / Published online: 13 July 2012 Ó Springer-Verlag 2012
Abstract The land snail family Odontostomidae has a poor fossil record, mainly from the middle Paleogene and early Neogene of Argentina. In this paper a new species of Odontostomidae from the Paleocene of Uruguay (Queguay Formation) is described. Bahiensis priscus n. sp. represents the first record of the genus Bahiensis Jousseaume 1877, and the oldest record for an Odontostomidae. The new species is characterized by a pupoid fusiform shell and an oval aperture with a single axial columellar fold. Present distribution of the genus indicates a tropical–subtropical environment, in high humidity rate areas. Keywords Fossil gastropods Odontostomidae Paleocene Paleosols Kurzfassung Die Landschnecken Familie Odontostomi¨ berlieferung, die in der dae besitzt eine spa¨rliche fossile U Hauptsache den Mittle Pala¨ogen bis Jung Neogen aus Argentinien umfasst. In dieser Arbeit eine neue Art der Familie Odontostomidae aus den Pala¨ogen aus Uruguay (Queguay Formation) wird beschrieben. Bahiensis priscus n. sp. erweist sich als dass a¨lteste Fund eines Odontostomidae. Die neue Art ist durch ein pupoid, spindelfo¨rmiges Geha¨use und eine ovale Mu¨ndung mit einer einzigen axialen Spindelfaltung charakterisiert. Die heutige Verbreitung der Gattung spricht fu¨r ein tropisches–subtropisches Ablagerungsraum, mit Ho¨hen Feuchtigkeit bzw. Humidita¨t werten.
F. Cabrera (&) S. Martı´nez Facultad de Ciencias, Universidad de la Repu´blica, Igua´ 4225, Montevideo, Uruguay e-mail:
[email protected];
[email protected]
Schlu¨sselwo¨rter Fossil gastropoden Odontostomidae Pala¨oza¨n Pala¨obo¨den
Introduction The superfamily Orthalicoidea is distributed in most of the Southern Hemisphere continents, but is especially abundant and diverse in the Neotropical region. It comprises the families Bulimulidae, Placostylidae, Odontostomidae, Amphibulimidae, Orthalicidae, and ?Megaspiridae (Breure et al. 2010, Breure and Romero 2012). The family Odontostomidae is restricted to South America, and is represented by at least 17 genera (Pilsbry 1902; Breure 1974; Richardson 1993; Simone 2006). It is characterized by an elongated spire with numerous whorls (more than six in several cases), and the presence of many folds in the aperture, which may be absent due to reduction (Pilsbry 1902; Breure 1974). The earliest record for the superfamily Orthalicoidea so far corresponds to several species assigned to Bulimulus and Itaborahia (Bulimulidae), from the Late Paleocene of the Sa˜o Jose´ de Itaboraı´ Basin (Rio de Janeiro, Brazil) (Brito 1967; Ferreira and Coelho 1971; Ferreira and Coelho 1989; Rodrigues and Da Fonseca 2007; Salvador and Simone 2011). Regarding Odontostomidae, there are until now only two fossil records: Plagiodontes sp. from the Middle Eocene–Oligocene, Sarmiento Formation (Miquel and Bellosi 2010), and ‘‘(?Cyclodontina) Plagiodontes dentatus’’ [sic] from the Upper Miocene, El Morterito Formation (Morton and Herbst 2007), both from Argentina. Cyclodontina dentata has been mentioned for the Upper Oligocene of Uruguay (Fray Bentos Formation) (Caorsi and Gon˜i 1958), cited later by Parodiz (1969) and Breure
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(1974), but it has been demonstrated that these gastropod specimens are recent ones drained into the older sediments (Figueiras and Broggi 1969; Martı´nez and Verde 1992). There is also a putative record from the Paleogene of Florida (USA) (Dall 1890; Breure 1974) that could be the only record of this family outside of South America. These fossils were described by Dall (1890), who assigned them to the genus Bulimulus. Subsequently, Breure (1974) considered that they should be placed under ?Anctus (Odontostomidae). Since they do not have any fold in the aperture, and the shell shape is similar to the representatives of Bulimulidae, these fossils should be included as originally proposed in this family and not in Odontostomidae. The aim of the present paper is to describe a new species of the genus Bahiensis Jousseaume 1877 (Odontostomidae) from the limestones of Queguay Formation from Uruguay (Late Paleocene–Early Eocene) from Quebracho locality Fig. 1 a Present distribution of the genus Bahiensis and b location map (Quebracho, Paysandu´–Uruguay)
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(Paysandu´, Uruguay) (Fig. 1), representing then the oldest record of the family, and the unique fossil record of the genus. Geological setting The Queguay Formation, or ‘‘Queguay limestones’’ in its informal name, are calcretes with an important fossiliferous content. The fossils, contained in their paleosols, consist of freshwater gastropods as Biomphalaria waltheri (Parodiz 1969) and Physa sp., terrestrial gastropods belonging to Bulimulus, Pupillidae, and Eoborus, ostracods, characeae oogons, hackberry endocarps (Celtis santosi), and ichnofossils (pupae and nests of Hymenoptera). The fossil assemblage indicates shallow palustrine facies and ponds of desiccation with periods of extreme aridity (Veroslavsky and Martı´nez 1996; Martı´nez et al. 1997; Martı´nez et al. 2001; Martı´nez and Veroslavsky 2004; Verde and Genise
Author's personal copy The oldest Odontostomidae (Mollusca: Gastropoda): Bahiensis priscus n. sp.
2007; To´falo and Morra´s 2009). Fossils contained in the paleosols were correlated with those of the lower strata of the Itaboraı´ Basin (e.g., by the presence of the genus Eoborus), suggesting a Late Paleocene age for Queguay Formation. These data are consistent with a continentalization verified in wells obtained from the continental platform, where a marked continentalization is shown by the disruption of the marine sedimentary strata in the Late Paleocene-Early Eocene, for about 15 million years (Martı´nez and Veroslavsky 2004; Daners and Guerstein 2004; To´falo and Pazos 2009; To´falo and Morra´s 2009). Taxonomy Pilsbry (1902) described 17 species of Bahiensis, some of them placed in other genera today. According to Simone (2006), the genus is composed by 12 living species (Simone 2006), most of them distributed only in Brazil; the exception is B. guarani that inhabits the Parana´ River Basin (Argentina, Brazil, and Paraguay) (Parodiz 1942b; Simone 2006) (Fig. 1). The literature about Bahiensis is scarce, being Pilsbry (1902) the main reference. Therefore, the relationships herein mentioned are supported only on morphological characters. Genus Bahiensis Jousseaume 1877. Type (by monotypy): Helix (Cochlogena) bahiensis Moricand 1833. Bahiensis priscus n. sp. Etymology
priscus from Latin: old, ancient.
Diagnosis Small, fusiform shell, with seven whorls, oval aperture with a visible axial columellar fold, outer lip thick and expanded. Description Small, fusiform shell, with seven slightly convex whorls, spire elongated. Narrow sutures, well defined. Sculpture composed of slight axial sigmoidal striae (3 per mm) more visible in the last whorl near the aperture; barely visible growth lines. Aperture oval, with a single inconspicuous axial columellar fold; thick outer lip expanded, arcuate and with a lower contact with the base. Last whorl represents 42 % of total shell length. Holotype: FCDPI 6474 (Fig. 2a–c). Height: 12.35 mm, maximum width: 4.45 mm, height of last whorl: 5.21 mm, height of aperture 3.9 mm, width: 2.9 mm. Other materials: FCDPI 6464 (Fig. 2d) and FCDPI 6469. Geographic and stratigraphic occurrence: Quebracho, Paysandu´ Department, Uruguay (Fig. 1), Queguay Formation, Late Paleocene-Early Eocene. The specimens are deposited in the paleontological collection of Facultad de Ciencias (Montevideo Uruguay)
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Facultad de Ciencias, Coleccio´n de Paleontologı´a–Invertebrados (FCDPI).
Discussion The specimens are included under the genus Bahiensis Jousseaume 1877 because of their thin, rather slender shell, composed of seven whorls, the apical ones with axial and spiral riblets, aperture rounded below, distal margin of the reflexed lip not separated from the whorl by grooves, and the presence of an axial columellar fold, all of these characters being distinctive of the genus (Pilsbry 1902). Regarding other genera of the family, the specimens do not belong to Cyclodontina Beck 1837, Clessinia Doering 1874, Spixia Pilsbry and Vanatta 1898, Plagiodontes Doering 1875–1876, or Odontostomus Beck 1837 because these genera have numerous teeth in the aperture, different in shape from those described for Bahiensis. Also, representatives of these genera are larger than these specimens. Plagiodontes and Odontostomus also differ in the shape of the spire, which is more globose than those of Bahiensis (Pilsbry 1902; Parodiz 1939; 1942a, b; 1944; 1948; Klappenbach and Olazarri 1973; Hylton Scott 1966; Breure 1974; Richardson 1993; Piza´ and Cazzaniga 2003). Species of Ventania Parodiz 1940 do not show any fold in the aperture margins, and the last whorl represents almost twothirds of the total length of the shell, being almost one-third of the total length of the specimens. Pilsbry (1902) did a revision of all the Odontostomidae described at that time. He was the first to put some order in the family, because even now, most of the species are mentioned under different genera (e.g., Odontostomus dentatus and Cyclodontina dentata). Pilsbry divided Bahiensis into two morphological groups, the second of them subdivided. The first one (bahiensis group) is composed by species with oblong, parallel-sided aperture, with the outer lip rather straightened; smooth surface with delicate axial striae; one (columellar) to four (parietal, columellar, upper and lower palatal) teeth, rarely with a developed suprapalatal fold. This group includes B. bahiensis (Moricand 1833), B. reevei (Deshayes 1851 in Fe´russac and Deshayes 1819–1851), B. occultus (Reeve 1849), B. ciaranus (Dohrn 1882), B. albofilosus (Dohrn 1883), and B. ringens (Dunker 1847). The second group (janeirensis group) consists of species with oval or oblong aperture, arcuate outer lip, and surface pitted, malleate, wrinkled or costulate. This group is secondary divided: the janeirensis–janeirensis subgroup is characterized by the absence of folds in the outer lip, and includes B. janeirensis (Sowerby 1833), B. miliola (d’ Orbigny 1834), B. rhodinostomus (d’ Orbigny 1835), B. guarani (d’ Orbigny 1834–47), and B. longulus (Pfeiffer 1859). The
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Fig. 2 a Ventral view of Bahiensis priscus n. sp.; b lateral view of B. priscus n. sp. (FCDP-I 6474) (scale: 5 mm); c columellar fold detail of B. priscus n. sp. (red arrow) (2 mm); d others specimens of B. priscus n. sp. (d FCDP-I 6464) (5 mm); e Bahiensis bahiensis
(Moricand, 1833) type species of the genus Bahiensis (front and lateral views) Cyclodontia [sic] (Bahiensis) bahiensis NMB 9612a (syntype) (10 mm); f Bahiensis guarani MACN 6654
janeirensis-punctatissimus group, with numerous folds or lamellae, comprises only B. punctatissimus (Lesson 1830). After Pilsbry’s review and until now, the taxonomy of Bahiensis remains unchanged. In a paper concerning the phylogeny of superfamily Orthalicoidea, Breure and Romero (2012) mention Bahiensis guarani as Cyclodontina guarani. Until a taxonomic revision of the family is done, we have chosen a conservative approach, including here this species into Bahiensis. Bahiensis priscus n. sp. is smaller than Bahiensis bahiensis, type species of the genus, and also differs in the shape of the aperture, being ovate in B. priscus n. sp. and oblong with parallel margins in B. bahiensis; the shape of the spire is similar in both (Fig. 2e). The new species from
the Paleocene of Uruguay has the characteristics of the janeirensis–janeirensis group, i.e., oval aperture and arcuate outer lip without folds (Fig. 3). Comparing with the other species of the genus, B. priscus n. sp. differs from B. janeirensis, B. rhodinostomus, and B. guarani (Fig. 2f) in the shape of the shell and number of whorls. These species have a conical-subfusiform shell, instead of the pupoid-fusiform shape of B. priscus n. sp.; B. janeirensis has eight whorls, and B. rhodinostomus and B. guarani have nine, while B. priscus n. sp. has seven whorls. The new species differs from B. longulus in the number of whorls (11 in B. longulus), and the shape of the shell and aperture (elongated in both cases). B. miliola has a parietal fold besides the
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Fig. 3 Sketch of the apertures of Bahiensis species, modified from original illustrations (see text for references). Scale bar: 5 mm. a–f bahiensis group: a Bahiensis bahiensis, b B. reevei, c B. ciaranus, d B. ringens, e B. occultus, f B. albofilosus; g– k janeirensis–janeirensis group: g B. janeirensis, h B. longulus, i B. guarani, j B. rhodinostomus, k B. miliola; l janeirensispunctatissimus group: B. punctatissimus; m Bahiensis priscus n. sp.
columellar lamellae and a strong ornamentation, instead of the single fold and slight striae of B. priscus n. sp. Modern species have the outer lip thinner than B. priscus n. sp. The outer lip is convex but straight in the modern species, being convex and arcuate in B. priscus (Figs. 2, 3), and the position of its contact with the base is lower than in the living species. Presently, the genus Bahiensis has a tropical–subtropical distribution, with its southern limit in northern Argentina (see ‘‘Introduction’’ and Fig. 1). Most of the species live in coastal damp forests, or rain forests: environments that share a high humidity rate. Consequently, it is not registered today in Uruguay, where these biomes are not present. According to this, the presence of Bahiensis in the fossil record might indicate tropical to subtropical climatic conditions for the Paleocene of Uruguay. This is consistent with the associated fossil assemblage and geological indicators as the calcrete-type limestones in which they are found (Martı´nez et al. 2001; Martı´nez and Veroslavsky 2004). Acknowledgments Mariano Verde collected the first specimens of this species. A. Batista, G. Roland, F. Montenegro, and N. Batalla gave field assistance, Alejandra Rojas helped with the images and language revision, Daniel de Amilivia helped with language, Eduard Sto¨ckli (Naturhistorisches Museum Basel, Basel, Switzerland) lent the type specimens of B. bahiensis, Sergio Miquel (Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia,’’ Buenos Aires, Argentina) allowed access to their collection, Peter Sprechmann translated the abstract to German, and Gabriela Cuezzo (Universidad Nacional Tucuma´n, Tucuma´n, Argentina) kindly answered some taxonomic questions. The reviewers Abraham Breure and Mathias Harzhauser made important suggestions to improve the paper.
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