First Record of Fossil Procyonid (Mammalia, Carnivora) from Uruguay T ∗ Leopoldo H
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Journal of South American Earth Sciences 92 (2019) 368–373 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames First record of fossil procyonid (Mammalia, Carnivora) from Uruguay T ∗ Leopoldo H. Soibelzona, , Andrés Rinderknechtb, Juliana Tarquinic, Raúl Ugalded,e a CONICET, Laboratorio de Morfología Evolutiva y Desarrollo (MORPHOS), División Paleontología de Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Paseo del Bosque s/n, B1900FWA, La Plata, Argentina b Museo Nacional de Historia Natural, 25 de Mayo 582, CC 399, Montevideo, 11000, Uruguay c Laboratorio de Paleontología de Vertebrados, Centro de Investigaciones Científicas y Transferencia de Tecnología a la Producción (CICYTTP-CONICET, Gob. de Entre Ríos, UADER), Materi y España s/n, E3105BWA, Diamante, Argentina d PEDECIBA Geociencias, Facultad de Ciencias, Universidad de la República, Iguá 4225, Montevideo, Uruguay e Escuela de Geología, Facultad de Ciencias, Universidad Mayor, Manuel Montt 367, Santiago, Chile ARTICLE INFO ABSTRACT Keywords: Procyonids were the first carnivorans that arrived from North America to South America during the late Camacho formation Miocene, before the full emergence of the Panama Isthmus. In South America, this family comprises five living Cyonasua genera (Bassaricyon, Nasua, Nasuella, Potos and Procyon) and two extinct genera (Cyonasua and Chapalmalania) Procyonidae that are recorded from the late Miocene to early Pleistocene. In this contribution, we report and describe spe- Late Miocene cimen MNHN 2965 assigned to Cyonasua sp., which represents the first record of Procyonidae from Uruguay. South America This specimen was collected at the coastal cliffs of Río de La Plata on the San José Department, in stratigraphic Uruguay levels that belong to the Camacho Formation (late Miocene). In addition, the presence of Cyonasua sp. in Camacho Fm. is in agreement with the substrate use and locomotor mode inferred for the genus (terrestrial and generalized, with some degree of climbing habits), and with what is known about the paleoenvironment of other formations were Cyonasua was found. 1. Introduction called “Cyonasua group”, then Forasiepi et al. (2014) demonstrated that they are a monophyletic group sharing a common ancestor with living Procyonids were the first eutherian carnivorans that arrived from procyonids. North America to South America during the late Miocene (Soibelzon There are ten formally named species of Cyonasua: C. argentina and Prevosti, 2007, 2012; Prevosti and Soibelzon, 2012), before the Ameghino (1885), C. brevirostris (Moreno and Mercerat, 1891), C. marine barrier, that separates North and South America, vanished circa longirostris (Rovereto, 1914), C. pascuali Linares (1981), C. groeberi 2.8 Ma (O'Dea et al., 2016). It is important to note that recent South Kraglievich and Reig (1954), C. lutaria (Cabrera, 1936), C. clausa American procyonids (Potos Geoffroy Saint-Hilaire and Cuvier, 1795; (Ameghino, 1904), C. robusta (Rovereto, 1914), C. argentinus Procyon Storr, 1780; Nasua Storr, 1780; Bassaricyon Allen, 1876; and (Burmeister, 1891) and C. meranii (Ameghino and Kraglievich, 1925), Nasuella Hollister, 1915) represent a second invasion from North or and a comprehensive taxonomic revision of these taxa is in progress. Central America during the latest Pleistocene-Holocene (Prevosti and One of the major problems is related to the nature of the holotypes, Soibelzon, 2012). since several of them are represented only by fragments of mandible or Five fossil procyonid genera were named in the past in South maxilla with very worn teeth and/or represented only by first to third America: Cyonasua Ameghino, 1885, Oligobunis, Burmeister (1891), premolars (P/p 1–3) without any systematic value (e.g., Cyonasua Pachynasua, Ameghino (1904), Brachynasua, Ameghino and Kraglievich clausa, C. robusta, C. argentinus). Additionally, most specimens in col- (1925), and Chapalmalania, Ameghino (1908). Of these only two are lections are catalogued as Cyonasua sp. or were assigned to species considered valid today (Cyonasua and Chapalmalania, Patterson and without explicit criteria (e.g., lower teeth assigned to a species which Pascual, 1972; Berman, 1994; Soibelzon and Prevosti, 2007; Soibelzon, holotype is a fragment of skull with some molars). Most specimens are 2011; Soibelzon and Prevosti, 2012; Forasiepi et al., 2014). Patterson classified based on where they were collected, at the same (or near) the and Pascual (1972) considered that both genera constitute a clade locality where the holotype of the referred species was collected. ∗ Corresponding author. Museo de La Plata, Paseo del Bosque s/nº, B1900FWA, La Plata, Buenos Aires, Argentina. E-mail addresses: [email protected] (L.H. Soibelzon), [email protected] (A. Rinderknecht), [email protected] (J. Tarquini), [email protected] (R. Ugalde). https://doi.org/10.1016/j.jsames.2019.03.024 Received 11 January 2019; Received in revised form 21 March 2019; Accepted 22 March 2019 Available online 27 March 2019 0895-9811/ © 2019 Published by Elsevier Ltd. L.H. Soibelzon, et al. Journal of South American Earth Sciences 92 (2019) 368–373 On the other hand, Chapalmalania includes two species: Ch. ortog- natha and Ch. altaefrontis whose holotypes are very well preserved. These two genera could be easily distinguished by their size and pre- dicted body mass, with Chapalmalania (88 kg; Prevosti and Forasiepi, 2018) much more robust than Cyonasua (range between 12.63 and 28.45 kg; Tarquini et al., 2018). The fossil record of Cyonasua is late Miocene to early Pleistocene (Huayquerian to Ensenadan Ages) while Chapalmalania is only recorded at the late Pliocene (Chapadmalalan to Marplatan Ages) Soibelzon and Prevosti (2012). Until now, the “Cyonasua group” was recorded in three countries: Argentina (late Miocene-early Pleistocene), Colombia (late Pliocene) and Venezuela (late Pliocene) (Soibelzon and Prevosti, 2012; Forasiepi et al., 2014). In this paper we describe the first record of Cyonasua sp. for the late Miocene (Camacho Formation) of Uruguay as a contribution to un- derstanding the natural history of these carnivorans in South America. Also we include some considerations on the paleoecology of Cyonasua. 2. Geological setting 2.1. Locality, age and geology The fossil-bearing strata are located in southwestern Uruguay (San José Department; 34° 34′ 38″S, 56° 58′ 32″W), in exposures of the coastal cliffs which are part of the Río de La Plata littoral platform (Fig. 1). Fig. 2. Stratigraphic section of the Puerto Arazatí exposures, with Cyonasua remains from the basal greenish claystone level. It also has two distinctive Crassostrea levels, at the base and at the upper portion. The right colored columns show different stratigraphic proposals for the units outcropping at the locality, with contrast on the upper contact of the Camacho Formation. (For interpretation of the references to color in this figure legend, the reader is re- ferred to the Web version of this article.) The lithostratigraphic units recognized from base to top are the Camacho (late Miocene), Raigón (Pliocene and Pleistocene) and Libertad (Pleistocene) formations (Bossi and Navarro, 1991; McDonald and Perea, 2002; Bossi et al., 2009; Tófalo et al., 2009; Perea et al., 2013). The first two units yield many vertebrate fossils, including a variety of cingulates, ground sloths, rodents and notoungulates (Vizcaíno et al., 2003; Rinderknecht et al., 2011, 2018; Perea et al., 2013). The remains described here were found in levels of the Camacho Formation (Fig. 2). This unit in Uruguay represents an extended late Miocene highstand eustatic event regionally known as the Paranean transgression or Paranean Sea (Sprechmann et al., 2000). In the San José Department, this includes facies related to a regressive phase, characterizing a shallow marine environment on a siliciclastic shelf. Terrestrial and marine vertebrates, in association with marine in- vertebrates and ichnofossils, comprise the main fossil assemblage Fig. 1. Study location and main places mentioned in the text. a) Broad and (Ubilla et al., 1990; Perea et al., 1996, 2013; Sprechmann et al., 2000; simplified map of the Uruguayan southwestern margin, showing yellow areas as Perea, 2005). The sediments of the Camacho Formation are formed by reference for Neogene sedimentary outcrops, the bigger coastal cities and the 87 86 greenish-grey friable claystones and mudstones that become greenish- Cantera Geymonat locality, where the Sr/ Sr ages were taken. b) Detail of the eastern San José Departament coast and the two principal localities for the brownish and slightly coarser toward the top. The lower boundary is Camacho Formation, Puerto Arazatí and Kiyú. (For interpretation of the re- not exposed. Former stratigraphic proposal (McDonald and Perea, ferences to color in this figure legend, the reader is referred to the Web version 2002; Tófalo et al., 2006; Bossi et al., 2009; Perea et al., 2013) sug- of this article.) gested different positions for the upper contact of the Camacho 369 L.H. Soibelzon, et al. Journal of South American Earth Sciences 92 (2019) 368–373 Fig. 3. Field attributes of the Puerto Arazatí outcrops of the Camacho Formation. a) Panoramic composite looking north of the coastal cliffs at the locality, showing the characteristic erosion patter in the upper levels, belonging to the Raigón Formation. The thin whitish, planar levels