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Molluscs from the Miocene Pebas Formation of Peruvian and Colombian Amazonia

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Molluscs from the Miocene Pebas Formation of Peruvian and Colombian Amazonia Molluscs from the Miocene Pebas Formation of Peruvian and Colombian Amazonia F.P. Wesselingh, with contributions by L.C. Anderson & D. Kadolsky Wesselingh, F.P. Molluscs from the Miocene Pebas Formation of Peruvian and Colombian Amazonia. Scripta Geologica, 133: 19-290, 363 fi gs., 1 table, Leiden, November 2006. Frank P. Wesselingh, Nationaal Natuurhistorisch Museum, Postbus 9517, 2300 RA Leiden, The Nether- lands and Biology Department, University of Turku, Turku SF20014, Finland (wesselingh@naturalis. nnm.nl); Lauri C. Anderson, Department of Geology and Geophysics, Louisiana State University, Baton Rouge, LA 70803, U.S.A. ([email protected]); D. Kadolsky, 66, Heathhurst Road, Sanderstead, South Croydon, Surrey CR2 OBA, England ([email protected]). Key words – Mollusca, systematics, Pebas Formation, Miocene, western Amazonia. The mollusc fauna of the Miocene Pebas Formation of Peruvian and Colombian Amazonia contains at least 158 mollusc species, 73 of which are introduced as new; 13 are described in open nomenclature. Four genera are introduced (the cochliopid genera Feliconcha and Glabertryonia, and the corbulid genera Pachy- rotunda and Concentricavalva) and a nomen novum is introduced for one genus (Longosoma). A neotype is designated for Liosoma glabra Conrad, 1874a. The Pebas fauna is taxonomically dominated by two fami- lies, viz. the Cochliopidae (86 species; 54%) and Corbulidae (23 species; 15%). The fauna can be character- ised as aquatic (155 species; 98%), endemic (114 species; 72%) and extinct (only four species are extant). Many of the families represented by a few species in the Pebas fauna include important ecological groups, such as indicators of marine infl uence (e.g., Nassariidae, one species), terrestrial settings (e.g., Acavidae, one species) and stagnant to marginally agitated freshwaters (e.g., Planorbidae, four species). Only seven species (4%) representing undisputedly elevated salinities were found, whereas 31 species (20%) are un- disputed restricted to freshwater biotopes. Only three (2%) terrestrial gastropod species are known. The Pebas system experienced profuse radiations of molluscs that led to an overwhelmingly endemic fauna, typical of a long-lived lake environment. Several extant genera, which nowadays live outside Amazonia, may have originated within the Pebas system. The stratigraphic continuity of species and line ages, at least for the late Early-early Late Miocene interval (c. 18-9 Ma), indicates that lakes continuously occupied the system and never were entirely replaced by rivers or the sea. The rare occurrence of marine taxa indi- cates that the system was at sealevel and occasionally experienced marine incursions. Contents Introduction .............................................................................................................................................................. 19 Material ....................................................................................................................................................................... 20 Systematic palaeontology ................................................................................................................................ 23 Discussion ................................................................................................................................................................ 279 Conclusions ............................................................................................................................................................ 282 Acknowledgements ........................................................................................................................................... 283 References ................................................................................................................................................................ 283 Introduction Nuttall (1990) reviewed the mollusc faunas of the Pebas Formation and other non- marine Neogene inland strata of northwestern South America. For the Miocene Pebas 20 Wesselingh. Molluscs from the Miocene Pebas Formation. Scripta Geol., 133 (2006) Formation of Peru and Colombia, and associated deposits in adjacent areas (including the ‘La Tagua Beds’ of southern Colombia, here considered part of the Pebas Forma- tion), Nuttall presented a fauna of slightly over 50 species. Extensive collecting during 1991 and 1996 by the present author added much material and many new species. Many of the species proved important for ecologic (Vonhof et al., 1998, 2003; Wesselingh et al., 2002, 2006c; Kaandorp et al., 2006), stratigraphic (Wesselingh et al., 2006a, b) and biogeo- graphic studies (Wesselingh & Macsotay, 2006a; Wesselingh & Salo, 2006). The aim of the present monograph is to document the mollusc species of the Pebas fauna. The Pe- bas fauna is defi ned here as the Miocene mollusc faunas from Amazonian and sub-An- dean basins, dominated in abundance and species numbers by corbulid bivalves and cochliopid gastropods. Some of the taxa from the Early Miocene ‘La Cira’ fauna (MZ1) of the Colombian Magdalena and Llanos Basins are assessed here as well in order to elucidate their relationship with the Pebas fauna. Material Institutional abbreviations – ANSP Department of Mollusca, Academy of Natural Sciences, Philadelphia, U.S.A. CAS Department of Palaeontology, California Academy of Sciences, San Fran- cisco, U.S.A. BMNH Department of Palaeontology, The Natural History Museum, London, England. INGEMMET Laboratorio de Paleontologia, Servicio Geologico, San Borja, Lima, Peru. LSU Department of Geology and Geophysics, Louisiana State University, Baton Rouge, U.S.A. NMB Naturhistorisches Museum Basel, Switzerland. RGM Division of Fossil Mollusca, Nationaal Natuurhistorisch Museum, Lei- den, The Netherlands (formerly Rijksmuseum voor Geologie en Mine- ralogie). RMNH Division of Mollusca, same institution, formerly Rijksmuseum voor Natuurlijke Historie. UNC Departamento de la Geologia, Universidad Nacional, Bogota, Colombia. Abbreviations – DL datum level of exposure (river level at date of collection in case of river exposures, surface of road in case of road exposure). fr. fragment. H height. Hap height of aperture, parallel to axis of the shell. L length. LV left valve. MZ mollusc zone (Wesselingh et al., 2006a). RV right valve. SA mean spiral angle (for adult specimens). SD semi-diameter (height of shell when viewed laterally). Wesselingh. Molluscs from the Miocene Pebas Formation. Scripta Geol., 133 (2006) 21 All measurements are in mm. Scale bars are in mm unless stated otherwise. The scales of illustrations of details should be inferred from the overall size of the shells. The mollusc biozonation scheme is that of Wesselingh et al. (2006a), which explains their stratigraphic distribution. Most of the shells were collected by F.P.W. in the Amazonas department, Colombia (1991) and the Peruvian department of Loreto (1991 and 1996; for localities see Fig. 1). The exposures are located in erosive bends of the Amazon River and its tributaries. No independent reference level or datum level can be assigned for such exposures, other than the relative position of a horizon with respect to the river level at the given date (which is given as DL). The section Santa Rosa de Pichana is a composite section, whose DL has been established by H. Vonhof and R. Kaandorp in 1998 (see Wesselingh et al., 2006c). The coordinates of the exposures are estimates from satellite imagery; those of a few localities (including Santa Rosa de Pichana) were checked later by colleagues using GPS and the original estimates were found to be only marginally incorrect. The newly collected material consists of over 3000 lots (Wesselingh et al., 2006a). Only material spe- cifi cally used in the descriptions is listed. Apart from the collected material, fossils were studied in several museums, including the BMNH and the ANSP. Genera of the Cochliopidae, Cerithioidea and Corbulidae are diagnosed and dis- cussed extensively. Diagnoses are more extensive for available species in these groups than for newly described taxa in order to allow for useful comparison. The species in species-rich genera are ordered by morphological similarity, in order to facilitate com- parison. The genera of other families are not treated in detail. Descriptions are based on material referred to as ‘material studied’ or ‘type material’. For available species, material published by Nuttall (1990), much of which was studied by the author during a visit to the BMNH in 1993, where at the time material also was available on loan pub- lished by de Greve (1938), has been used to add to the descriptions. Type specimen data of almost all existing species has been given in Nuttall (1990). Some 80 cochliopid spec- imens were subjected to a SEM study of apical details. The general discussion on Coch- liopidae and the section on Longosoma (three species) are by F.P.W. and D. Kadolsky, whereas L. Anderson and F.P.W. described Corbula cotuhensis. Systematic palaeontology Class Gastropoda Family Neritidae Lamarck, 1809 Genus Neritina Lamarck, 1816 Type species – Nerita pulligera Linné, 1766, Recent, Indo-Pacifi c. Remarks – Wesselingh (2003) discussed the (sub)generic attribution of the Peba- sian neritines, but could not resolve this matter. One of the possible candidates, Neri- tina (Fluvinerita), is amongst others characterised by the lack of an opercular peg (Nut- tall, 1990). Pebasian neritine opercula studied by Nuttall (1990) and Wesselingh
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