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(Aves, Cathartidae) from the Late Pleistocene of Uruguay

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(Aves, Cathartidae) from the Late Pleistocene of Uruguay Journal of South American Earth Sciences 107 (2021) 102946 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames First report of large cathartids (Aves, Cathartidae) from the late Pleistocene of Uruguay W.W. Jones a,d,*, A. Rinderknecht a, R.I. Vezzosi b,c,d, F. Montenegro a,e, M. Ubilla d,e a Museo Nacional de Historia Natural, 25 de Mayo 582, CP 11000, Montevideo, Uruguay b Laboratorio de Paleontología de Vertebrados, Centro de Investigaciones Científicas y Transferencia de Tecnología a la Produccion´ (CONICET, Gob. E.R., UADER), Materi y Espana,~ Diamante, E3105BWA, Argentina c Facultad de Ciencia y Tecnología, Universidad Autonoma´ de Entre Ríos, Ruta Provincial 11 km 10,5, Oro Verde, E3100XAD, Entre Ríos, Argentina d Programa de Desarrollo de las Ciencias Basicas´ (PEDECIBA – UDELAR), Uruguay e Departamento de Paleontología, Facultad de Ciencias (UdelaR), Igua´ 4225, CP 11400, Montevideo, Uruguay ARTICLE INFO ABSTRACT Keywords: The fossil record of South American cathartids, with few exceptions, is largely restricted to Late Pleistocene and New World vultures Holocene sites. This contribution provides for the first time fossil records of cathartids from Uruguay. The Scavenger birds specimens reported here include an; almost complete fibula, an incomplete furcula, and the distal end of a Guild tibiotarsus. The; firsttwo specimens came from Late Pleistocene beds at two localities of from Northern Uruguay, Late Pleistocene and the last one comes from Late Pleistocene-Early Holocene beds from South-western Uruguay. All the speci­ Uruguay mens were associated with several megafaunal fossil remains. The systematic assignation and paleobiological implications of these scavenger birds are here discussed. 1. Introduction America and the Antilles (see Cuello, 1988 and references therein; Suarez,´ 2020), most notably the fossil cathartid richness from Rancho La New World vultures (Family Cathartidae; for a systematic proposal Brea, in California, United States (4 genera and species; see Miller, 1909, see Remsen et al., 2020 and Winkler et al., 2020) are scavenger birds 1911, 1925; Howard, 1974; Emslie, 1988b). In summary, the Cathar­ widely distributed throughout the America. Cathartidae is composed of tidae is one of the best documented avian families in the Quaternary of seven extant species: the large Andean and California condors Vultur the Americas. Nevertheless, it is remarkable that there are practically no gryphus Linnaeus, 1758 and Gymnogyps californianus (Shaw, 1797), anatomical, systematic and phylogenetic works that allow for the respectively; the intermediate sized King Vulture Sarcoramphus papa elucidation of the taxonomic richness of New World vultures. (Linnaeus, 1758) and the smaller vultures that include the Turkey This contribution provides the first report of cathartids from Vulture Cathartes aura (Linnaeus, 1758); the Greater and Lesser Uruguay. The materials come from Late Pleistocene localities, and are Yellow-headed Vultures Cathartes melambrotus Wetmore (1964) and associated with Lujanian mammal remains. The systematic assignment Cathartes burrovianus Cassin (1845) respectively, and the Black Vulture and a brief discussion of the paleobiological implications of these find­ Coragyps atratus (Bechstein, 1793). ings are provided. Extant cathartids in South America were reported repeatedly in Late Pleistocene and Holocene sites from Argentina, Brazil, Ecuador and Peru 2. Geographic and geological provenance (Brodkorb, 1964, p. 257; Campbell, 1976, 1979; Alvarenga, 1998; Tonni and Noriega, 1998; Tambussi and Noriega, 1999; Noriega and Areta, The materials studied in this work include three fossil (a tibiotarsus, 2005). Thus far, four extinct genera and fivespecies have been described fibula, and furcula) from three different localities (Fig. 1). from Peruvian to Argentinean Pleistocene outcrops (Campbell, 1979; The tibiotarsus (CB 109) was collected by Rolando Bianchi from Tambussi and Noriega, 1999; Alvarenga and Olson, 2004; Noriega and Prestes glen (rural establishment El Bravío, Department of Soriano, SW Tonni, 2007; Alvarenga et al., 2008; Agnolin et al., 2017). This record is Uruguay) 7.9 km from Dolores city. These levels were assigned to the extended if one considers the discoveries from North and Central Dolores Formation, which is a late Pleistocene-early Holocene age unit * Corresponding author. Museo Nacional de Historia Natural, 25 de Mayo 582, CP 11000, Montevideo, Uruguay. E-mail address: [email protected] (W.W. Jones). https://doi.org/10.1016/j.jsames.2020.102946 Received 31 July 2020; Received in revised form 5 October 2020; Accepted 5 October 2020 Available online 10 October 2020 0895-9811/© 2020 Elsevier Ltd. All rights reserved. W.W. Jones et al. Journal of South American Earth Sciences 107 (2021) 102946 County Natural History Museum; MHNT: Museu de Historia Natural de Taubate;´ MLP OR: Museo de La Plata; MNHN: Museo Nacional de His­ toria Natural de Uruguay; USNM: Smithsonian Natural History Museum. 3.2. Comparative sample We follow the phylogenetic arrangement among the order Cathar­ tiformes given by Emslie (1988a). The “condor group” is recognized by the cited work and previous authors termed it Vulturinae, after Camp­ bell (1979). Nevertheless, Johnson et al. (2016) proposed Sarcoramphus to be well-nested among condors (sensu Brito, 2008). However, we taken into account Agnolin et al. (2017) recommendations that discard this arrangement until new evidence sustaining such a proposal becomes available. We considered New World Vultures as within Order Cathar­ tiformes, rather than as Cathartidae within Accipitriformes, following Remsen et al. (2020). Until now, the unique phylogenetic proposal of family Cathartidae based on morphological analysis was performed by Emslie (1988a), which included Teratornithidae in the order Ciconiiformes and consid­ ered this family to be a sister group of Cathartidae. Moreover, Ter­ atornithidae has been traditionally considered to be closely related with Cathartidae (Miller, 1909, 1925). In fact, Brodkorb (1964) has consid­ Fig. 1. Map of Uruguay showing the location of the findings:Sopas Formation: ered teratornithids as a subfamily of Cathartidae, but the treatment of A) Sopas Creek (Department of Salto), B) Malo Creek (Department of this taxon as an independent family has prevailed (see De Mendoza and ´ Tacuarembo); Dolores Formation: C) Prestes glen (Department of Soriano). Picasso, 2019; Suarez,´ 2020). The presence of putative teratornithids among late Pleistocene avifauna of South America has been previously (see Nami, 2020; Ubilla et al., 2018 and references therein). The Prestes mentioned (Campbell, 1976; Campbell and Tonni, 1983; Agnolin, 2016; glen outcrop yielded fossils of Quaternary megafauna such as ground Cenizo et al., 2016). However, the taxonomic identity of these remains sloths (Lestodon armatus), notoungulates (Toxodon platensis), camelids has not yet been reviewed. For all of these reasons, we compared the (Hemiauchenia paradoxa) and gliptodonts (Gliptodontidae sp.), among fossil remains presented in this work not only with Cathartidae but also others. with Teratornis merriami, the most complete teratornithid species of the The geographic and stratigraphic provenance for the fibula(FC-DPV- Late Pleistocene in North America. 2673; discovered by one of the authors A.R.) is Sopas creek, (Salto The osteological material consulted for the extant Cathartidae spe­ Department, NW Uruguay), Sopas Formation. The sediments of this cies includes the following: Cathartes aura MNHN 7005, 7012, MHNT Formation were assigned to the late Pleistocene (Ubilla et al., 2004). 47, 794, USNM 562524, 346582; C. burrovianus MHNT 714; This locality in particular was dated with the OSL method in 43.5 ± 3.6 C. melambrotus MHNT 1213; Coragyps atratus MNHN 6292, MHNT 730, ka (basal level); 30.6±5.4 ka(paleocave infilling sediments; see Sup­ 985, 10032, 10025, 10040; Gymnogyps californianus LACM 10181, plementary Material and Ubilla et al., 2016). USNM 3369, 346582; Sarcoramphus papa CFA-OR-1125, MHNT 1787, The geographic and stratigraphic provenance for the furcula (FC- 804, 775, 903, USNM 559318; and Vultur gryphus MHNT 591, 124303, DPV-2881; discovered by one of the authors A.R.) is Malo creek, MLP OR 367, USNM 623236, 430210. (Tacuarembo´ Department, N Uruguay), Sopas Formation, Late Pleisto­ The fossil material consulted includes Teratornis merriami LACM 14 cene. This locality was dated with the AMS C method (ages: 33.6 ± 0.7 4426, B1270; and Vultur gryphus (Talara Seep Deposits) ROM 16801, ka B.P. (cal 36.1–39.4 ka) to 39.9 ± 1.1 (cal 42.0–45.4 kya) and the OSL 16803, 16805. The fossil material of Breagyps clarki, Gymnogyps ages: 58.3 ± 7.4 to 32.8 ± 1.9 ka (see Supplementary Material and Ubilla howardae, and Geronogyps reliquus were accessed by descriptions and et al., 2016). The furcula was found associated with the level of AMS 14C illustrations previously published (Howard, 1974; Campbell, 1979). cal 36.1 39.4 ka. The OSL and fresh-water mollusk shells AMS 14C ages The fossil materials here studied are hosted in the Coleccion´ Pale­ from the Sopas outcrops indicate a relationship with the MIS-3 (Ubilla ontologica´ (Vertebrados Fosiles),´ Facultad de Ciencias, Universidad de et al., 2016). la República, Montevideo, Uruguay (FC-DPV 2673, 2881) and the The fossil record of the Sopas Formation includes a large taxonomic Rolando Bianchi paleontological collection, Dolores, Department
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