Mollusca, Gastropoda) Brazil
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October 2003 CainozoicResearch, 2(l-2)(2002), pp. 117-128, Neritidae (Mollusca, Gastropoda) from the Miocene of western Amazonia (Peru, Colombia and Brazil) Frank+P. Wesselingh NationaalNatuurhistorisch Museum (Naturalis), P.O. Box 951 7, NL-2300 RA Leiden, the Netherlands; e-mail: [email protected]; and, University of Turku Biodiversity Centre, Faculty ofBiology, University of Turku, SF-200I4 Turku. Finland Received 25 January2003, revised version accepted 24 March 2003 Six neritid species are recorded from the Miocene (PebasFormation) of western Amazonia (Peru, Colombiaand Brazil), two of which, The Neritina (?) patricknuttalli and N. (?) elephantina, are described as new. subgeneric status of Pebasianneritids remains unresolved. Palaeoecological and palaeobiogeographicalimplications are briefly discussed. KEY WORDS:: South America, Amazonia, Miocene, Neritidae, Neritina, new species. Introduction examine relationships with other Recent and fossil neo- tropical neritids, and to discuss palaeoecological and In the well-preserved Miocene molluscan faunas of the palaeobiogeographic implications. Because of the uncertain Pebas Formation in western Amazonia, specimens of subgeneric status of the six species of Neritina known to Neritina are common. In his seminal review of South date from the Pebas Formation and of its close relatives American non-marine molluscan Nuttall the informal ‘Pebasian is used faunas, (1990) (see below), name Neritina’ considered all Miocene species ofNeritina from the west- here to characterise this group. ern Amazonia to be conspecific and he used the name ?N. ortoni Conrad, 1871 for them. With the exception of a single neritid from the Cantaure Formation (Miocene) of Systematic palaeontology northernVenezuela, Nuttall was unableto link the Pebasian fossil The Pebasian Abbreviations — denote the of material neritids to any other or extant taxa. To repositories neritids are characterised by their commonly velatiform referred to in the text, the following abbreviations are used: outline (i.e., expanded inner lip callus, basal portion of shell’s body whorl in a plane strongly inclined to the axis, and flaring outer lip) and by the absence of an opercular BMNH The Natural History Museum, London (formerly peg. The latter feature is shared with extant N. (Fluvinerita) British Museum [Natural History]); tenebricosa (Adams, 1851) from Jamaica, but Nuttall CAS CaliforniaAcademy ofSciences, San Francisco; (1990) found that species to differ from the Pebasian GG collections of the Department of Palaeontology neritids by its thick, spirally striated, non-velatiform shell, (BMNH; see Nuttall, 1990); and dismissed it as a possible close relative of Pebasian 1NGEMMET Geological Survey, San Borja, Lima; Neritina. A study of newly collected material from the NYSM New York State Museum, Syracuse; Pebas Formation, and a re-examinationof museum speci- PIMUZ Palaontologisches Institut und Museum der from ofnorthwest South America Universitat mens Neogene deposits Zurich; have now shown Pebasian Neritina to comprise several RGM Nationaal Natuurhistorisch Museum, Leiden species. These species are most easily, but not exclusively, (formerly Rijksmuseum van Geologic en Miner- distinguished by their colourpatterns, although this feature alogie), Department of Palaeontology, Division is notorious for its intraspecific variation (see e.g., Will- ofCainozoic Mollusca for for Nationaal Natuurhistorisch mann, 1981 Theodoxus; Costa et al., 2001 RMNH Museum, Leiden Neritina). (formerly Rijksmuseum voor Natuurlijke Histo- the to for of The aim of present paper is provide diagnoses ric), Department Evertebrates, Malacology differentiationof Miocenewestern Amazonian neritids, to Division. - 118 - Stratigraphic age estimates are adapted from Hoorn (1993); having smaller, higher-spired shells, with more whorls, a data are listed in 1, while the material in dental architecture and of locality Appendix non-flaring outer lip, presence studied, with the exception of type specimens, is listed in a dark colour band subsuturally on most specimens. Nuttall 2. Appendix (1990) discussed and rejected a close relationship between Pebasian Neritina and extant Neritina zebra (Bruguiere, 1792). Neritilia succinea (Recluz, 1841 (extant, Greater Superfamily Neritoidea Lamarck, 1809 Antilles and central America) has a ‘subpatelliform’ outline Neritidae 1809 Family Lamarck, (Russell, 1941, p. 398) in common with the Pebasian Genus Neritina Lamarck, 1816 of it is small width species Neritina. However, very (shell < 6 mm) and the colour is deeply impregnated into the The Pebasian neritids characterised velatiform are by a shell, while in the Pebasian species colours form part ofthe shell form and the absence ofan opercular peg. Using these outer shell layers only, and easily come off. The Pebasian features, the following neotropical species are attributedto neritids differ from other extant neotropical neritids in the ‘Pebasian Neritina group’: having a low spire (see e.g., Russell, 1941). The range of morphological variation resembles that of the Eurasian * Miocene, western Amazonia: genus Theodoxus de Montfort, 1810. It is only by virtue of Neritina(?) ortoni Conrad, 1871, A. (?)puncta Etheridge, the disjunct temporal and geographic distribution, as well 1879 N. (?) etheridgei Roxo, 1924, N. (?) roxoi de Greve, as the tropical distribution of the Pebasian species of N. and N. distributionof Theo- 1938, (?)patricknuttalli n. sp., (?) elephantina n. Neritina, vs a temperate-subtropical sp. doxus, that these two taxa are treated as distinct forthe time being. There is no living species of the ‘Pebasian Neritina * Miocene, northern Venezuela: group’ to test this. The shelloutline ofseveral of the Peba- N. aff. woodwardiGuppy, 1866 sian neritids superficially resembles that of the Caribbean Neritinapunctulata Lamarck, 1816 (Figure 21). However, * Ecuador: the of close Miocene, presence an opercular peg argues against a N. pacchiana Palmer in Liddle & Palmer, 1941 relationship between these two. Isolated opercula (see Nuttall, 1990, fig. 26a, b) are Three other dominicana species, Neritina Costa et al, common in the Pebas Formation, but they cannot be attrib- 2001 and N.jungi Costa etai, 2001 (Miocene, Dominican uted to any particular species with certainty. All of these Republic) and#, sp. Woodring, 1973 (Eocene, Panama) opercula, however, lack a peg, and it is therefore safe to the ‘Pebasian with that do indeed are assigned to Neritina group’ some assume they belong to the Pebasian species hesitation (see Discussion). documentedbelow. Initially, I placed Pebasian species ofNeritina illthe sub- — Small to width low- Diagnosis large (shell 4-22 mm), genus Fluvinerita Pilsbry, 1932 (extant, freshwater, Ja- ovate to spired, globose, commonly shouldered neritid; maica), on the basis of the absence of an opercular peg. low-spired (velatiform) species commonly involute, with a However, a recent discussion with H.K. Mienis (Hebrew hooked suture viewed from above; ovate species with University, Tel Aviv) has convinced me that several other nucleus and characters prominent erect a suture describing a spiral displayed by the Jamaicantaxon (<e.g., absence when viewed from subsutural of denticles columellar above; ramp convex to (more on lip, a comparatively poorly commonly) concave; growth lines strongly prosocline; developed inner lip callus, a higher spire and a rather shells with a wide range of colour patterns, including efface convex whorl profile) are incompatible with the Pebasian white and colour and the to zigzag punctate; patterns superficial; neritids, previous subgeneric assignment was diffuse spiral striae visible on abraded (portions of) speci- abandoned. subdued and/oraxial ribs but characteris- mens; spiral rare, tic of a single species; very fine, minute, slightly irregular spiral cords on very well-preserved specimens of most Neritina (?) ortoni Conrad, 1871 inner species; a flaring outer lip on most larger species; lip Figures 1-7 callus well developed to prominent, often thickened; callus in the entire basal - expanding some species over shell; 1869 Neritina pupa (Linne) Gabb, p. 197, pi. 16, fig. 2. * 1871 Neritina ortoni operculum lacking peg. Conrad, p. 195, pi. 10, figs 5,11. 1871 Neritina - ortoni Conrad Woodward, p. 103, pi. 5, fig. 2a, b. — of the have Differentiation Species genus Puperita 1878 Neritina - ortoni Conrad Boettger, p. 428. obscure spiral ribs (Costa et al., 2001), but are compara- 1938 Neritina (Vitta) amazonensis de Greve, p. 62, pi. 5, figs tively high-spired and lack the prominent columellar area 4-6, 8. of the of Neritina discussed herein. The typical species (?) 1938 Neritina Roxo - (Vitta) etheridgei de Greve, p. 66, pi. 5, of Neritina 1917 group fulgopicta Maury, (Miocene, figs 1-3, 7, 9. Dominican and N. 1966 ?Neritina de Greve — Republic) virginea (Linne, 1758) (ex- amazonensis Willard,p. 66, pi. 62, tant, Caribbean) differs from the Pebasian neritids in fig. 3. - 119- All specimens illustrated, with the exception of Figure 21, are from the Miocene Pebas Formation; for locality data see Appendix 1 Neritina (?) ortoni Conrad, 1871 - 1 lectotype, NYSM 9277 (J. Hauxwell Colin), ‘Pichua’ (probably Santa Rosa de Pichana), Loreto department, Peru; 2 - paralectotype, NYSM 9276, same locality; 3 - ROM 456 092, Santa Rosa de Pichana, Loreto department, Peru; 4 - RGM 456 100, Santa Elena III, Loreto department, Peru (collected from surface); 5 - RGM 456 134, C14 Hoom Colin, Santa Julia, Loreto department,Peru; - 6