Journal of South American Earth Sciences 107 (2021) 102946

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Journal of South American Earth Sciences

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First report of large cathartids (Aves, Cathartidae) from the late 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 record of South American cathartids, with few exceptions, is largely restricted to and New World sites. This contribution provides for the first time fossil records of cathartids from Uruguay. The Scavenger 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 , 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 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 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 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 “ 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 , 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 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 of Quaternary 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 merriami, the most complete teratornithid species of the The geographic and stratigraphic provenance for the fibula(FC-DPV- Late Pleistocene in . 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 of diversity of vertebrates, mostly represented by as a dominant Soriano, Uruguay (CB-109). group (ca. 25 families and more than 50 species). The accompanying The anatomical nomenclature follows Baumel and Witmer (1993) mammal fauna of the studied material encompasses mega and large and additional terms proposed by Livezey and Zusi (2006). The mea­ extinct xenarthrans, immigrant and autochthonous ungulates, small to surements were taken with a digital caliper with an accuracy of 0.01 large rodents and a variety of carnivores, among others (Ubilla et al., mm. For comparisons, we used the published measurements of Fisher 2004, 2011; 2016; Gasparini et al., 2013; Manzuetti et al., 2018, 2020; (1946, 1947), Campbell (1979), and Howard (1974). Morosi and Ubilla, 2019). 4. Results 3. Materials and methods 4.1. Systematic paleontology 3.1. Institutional abbreviations Remsen et al., 2020. CFA-OR: Felix´ de Azara Ornithology Collection, Fundacion´ de His­ Cathartidae Lafresnaye, 1839 (sensu Remsen et al., 2020). toria Natural “Fundacion´ Felix´ de Azara”, Buenos Aires, Argentina; CB: Cathartidae indet. Rolando Bianchi, Paleontological Collection, Dolores city, Uruguay; FC- DPV: Coleccion´ Paleontologica´ (Vertebrados Fosiles)´ of Facultad de 4.1.1. Referred material Ciencias, Universidad de la República, Uruguay; LACM: Los Angeles FC-DPV 2673 almost complete right fibulawith missing distal spine

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However, Jollie (1976) and Fisher (1946) noted differences in the procesus interclavicularis that forms the junction of the two furcular rami in these taxa. This small process tapers and is anteroventrally pointed in Cathartidae; it is notoriously less tapered in T. merriami (Miller, 1925; Fisher, 1946; Jollie, 1976). Moreover, the furcular ramus in T. merriami is less robust than in all living cathartid species (Fig. 5). In cathartids, the anterior and internal furcular surfaces are smooth in contrast to the sharp edges observed in Accipitridae (Fisher, 1946). Both furcular traits allow to assign the material studied here to Cathartidae. In terms of size (Table 2), FC-DPV-2881 can be included in the inferior limit of the Vultur gryphus range size that is represented by condor fe­ males. However, its generic level assignation is not clear due to its poor preservation.

4.3. Cathartidae indet

4.3.1. Referred material CB-109 distal portion of the right tibiotarsus (Fig. 6).

4.3.2. Comparative osteology The distal width is narrower than in falcons and accipitrids, and corresponds with the observed in cathartids (Lydekker, 1891). The Fig. 2. Fibula of Cathartidae indet. (FC-DPV 2673 ) in lateral (A), posterior (B), lower margin of the supratendinal bridge curves upwards, resulting in a proximal (C), anterior (D) and medial (E) views. Scale bar equals 1 cm. large, unevenly shaped opening of the canalis extensorius (as is observed in all of the living and fossil cathartids, see Howard, 1974). The medial (Fig. 2). axis of this canal is aligned with the medial shaft, like in Gymnogyps californianus and Sarcoramphus papa. The same canal differs notoriously 4.1.2. Comparative osteology in teratornitid species, with a more expanded shape and consequently a ´ Teratornis merriami is differentiated from cathartids in having prox­ diffused lateral rim (see Miller, 1925, plate 4; Suarez and Olson, 2009, ´ imal articular surface of the fibularhead with straighter rims; the lateral Fig. 8; Suarez, 2020, Fig. 5A; Fig. 7, this work). Jollie (1976, Fig. 180) rim continues in a notoriously steeper angle not observed in cathartids shows the occurrence of a fovea or lateral canaliculus to the canalis (the insertion of ligamentum tibiofibulare cranialis is located here; extensorius in cathartid species, which is more conspicuous in Vultur Fig. 3). In the lateral view Vultur gryphus, Gymnogyps californianus, and gryphus (fovea ligamenti collateralis lateralis sensu Livezey and Zusi, Coragyps atratus shows a prominent proximal process that points pos­ 2006; Brito, 2008; Alvarenga et al., 2008). This fovea is located proxi­ teriorly and that it is absent in Cathartes spp. and T. merriami. Although mally near to the condylus lateralis in the material studied here (as in this process is distally broken in the material studied here, the base is Gymnogyps californianus, more distally located in V. gryphus and S. papa. clearly differentiated (Figs. 2E and 3B). The fibular head in T. merriami The tuberositas retinaculi extensoris are verys similar in shape and exhibits the major convexity of the proximal rim, whereas in FC-DPV location among cathartids and teratornithids, and a great intraspecific 2673 this rim is slightly convex, as in V. gryphus and G. californianus; variation is also observed. In CB 109, the tuberositas proximalis reti­ in C. atratus and Cathartes spps., this rim is nearly straight (Fig. 3B). naculus extensoris is partially broken, it was relatively large and tear­ According to the ligamentous insertions of FC-DPV 2673, the area of drop shaped, similar to cathartid and T. merriami condition. The ligamentum tibiofibulare cranialis is partially broken. The insertion of tuberositas distalis is represented as a pronounced tuberculum in all the ligamentum collateralis lateralis has a subtriangular shape in all cathartids (including CB 109) and T. merriami. This tubercle in CB 109 is living cathartids whereas it has a subelliptical contour in T. merriami. roughly subtriangular in contour, as occurs in V. gryphus and S. papa. The insertion of the musculus plopliteus has a subcircular shape and is This tuberculum is more prominent and quadrangular in shape in caudally oriented with respect to the fibular head in the fossil studied T. merriami and G. californianus. The lateral canaliculus is laterally here and the other cathartids; this impression is notoriously elongated in located with respect to this tuberculum in T. merriami, G. californianus; it T. merriami and is more caudally positioned. The ansa m. iliofibularisis has a distal location with respect to this tuberculum in CB 109, insinuated in the lateral facet of FC-DPV 2637 and follows an oblique V. gryphus and S. papa. The condylus lateralis is laterally oriented as is orientation, as in cathartids and teratornithids. Gross shape and size of observed in Pleistocene Vultur; it is not flaredas V. gyrphus, nor aligned the proximal articular facet of the fibulareported is comparable to that with the longitudinal axis of the diaphysis as in G. californianus and of G. californianus (Fig. 3 and Table 1). S. papa. The condylus medialis is medially flaredas in Vultur (less abrupt flaring occurs in Gymnogyps and Sarcoramphus). T. merriami presents both cartiliginis trochlea very low with respect to cathartids, with the 4.2. Cathartidae indet exception of the living S. papa (Miller, 1909, 1925, see Fig. 7). The incisura intercondylaris in T. merriami is transversely wider (at least 4.2.1. Referred material proximally sensu Suarez´ and Olson, 2009) than in cathartids (including FC-DPV 2881 incomplete furcula with an almost complete right the CB 109). Compared with fossil cathartids from Talara Seeps, Peru clavicle, apophysis furculae and an almost incomplete left clavicle (Campbell, 1979), Gymnogyps howardae and Geronogyps reliquus, the (Fig. 4). fossil tibiotarsus described here shows similar size (see Table 3). CB 109 and G. reliquus are similar in the location of the canaliculus lateralis near 4.2.2. Comparative osteology of lateral condylus, and in the alignment of the sulcus extensorius with Jollie (1976) remarks that the extremitas omalis claviculae has the shaft longitudinal axis. On the other hand, G. reliquus clearly differs several diagnostic traits observed in the living species of cathartids and from CB 109 in having a deep internal ligamental attachment. Teratornis merriami. The preservation of the fossil material The comparison of the tibiotarsus size and the shape of sulcus (FC-DPV-2881) does not allow the observation of the omal extremity. extensorius and condylae clearly discard affinities with the small

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Fig. 3. Comparative views of living cathartids and extinct Teratornis merriami fibulae.A) Proximal articular surfaces (not to scale). B) Lateral views (scale bar equals 1 cm). Abbreviations: Ansa m. iliofibularis = Ansa musculi iliofibularis, cap. fib. = caput fibulae, Impress. lig. tibiofibularis cranialis = Impressio ligamenti tibio­ fibularis cranialis, Imp. m. popliteus = Impressio musculi popliteus.

Table 1 Fibular measurements (in mm) of extant and fossil species (†).

Specimen Cathartidae sp. Indet. FC- Teratornis merriami† Vultur Gymnogyps californianus LACM Sarcoramphus papa CFA-OR DPV 2673 LACMB4426 gryphus 101811 1125 N = 2

Fibular head length 16.50 est. 20.13 19.86 17.9 12.71 Fibular head-tuberculum m. 49.68 56.56 54.84 52.47 37.52 iliofibularis length

Abbreviation: est. (estimation). cathartid Pleistovultur nevesi Alvarenga et al., 2008. The tibiotarsus of 5. Discussion Breagyps clarki has a mean transverse width of distal end of tibiotarsus similar to CB 109 (Howard, 1974, Table 9: mean = 24.4 mm). However, 5.1. Taxonomic remarks it differs from B. clarki in the pronounced internal flaringof the condylus medialis, the absence of a medial projection of this condylus and the Clearly, the three Uruguayan fossils studied here yield osteological presence of a lateral projection in condylus lateralis (Howard, 1974, features that allow them to be assigned to the family Cathartidae (e.g. Fig. 9). The distal end width indicates that the CB 109 is smaller and out the shape and robustness of the furcular rami and disposition of the of the range size of Vultur gryphus. In size terms, the fossil tibiotarsus processus interclavicularis; the shape of the fibular head and ligamen­ presented here has similar dimensions to the extant Gymnogyps cal­ tous attachments; a narrow distal width of the tibiotarsus; and the width ifornianus (Table 3). and lateral rim of the sulcus extensorius). Some osteological features and size measurements of tibiotarsus CB 109 and fibulaFC-DPV 2673 suggest affinitieswith the living California

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Fig. 4. Furcula of Cathartidae indet. (FC-DPV 2881) in anterior (A), lateral (B) and posterior (C) views. Scale bar equals 1 cm.

Fig. 5. Comparative anterior views of furcula: A) Teratornis merriami, B) FC-DPV 2881, C) Vultur gryphus, D) Gymnogyps californianus, E) Sarcoramphus papa. Ab­ breviations: proc. interclav. = procesus interclavicularis. Scale bar equals 1 cm.

Table 2 Clavicular symphysis width of fossil and extant Cathartidae (in mm).

Cathartidae Vultur Vultur gryphus Gymnogyps Sarcoramphus sp. Indet. FC- gryphus USNM430210 californianus papa USNM DPV 2881 MHNT USNM 346582 559318 591

23.71 25.97 27.8 19.7 15.26

Condor Gymnogyps californianus, a living relict cathartid species from North America (Fisher, 1947; Emslie, 1987). However, the fossil record shows a larger distribution of this during the late Pleistocene. In North America and Cuba G. kofordi Emslie, 1988a, and G. varonai (Arredondo, 1971) respectively, and G. howardae Campbell, 1979 for Northern South America. This record shows the large dispersion of this taxon and allows us to contemplate the possibility of assigning it to the fibula FC-DPV 2881 and tibiotarsus CBF 109 to this genus, even assigning it to a new species of Gymnogyps. However, it is clear that more osteological elements are required to perform a robust taxonomic designation. Based only on a rough calculation of proportional dimensions we cannot rule out that the three fossils studied here could represent a single taxon. However, the different provenances and ages make this claim tenuous.

5.2. Paleoecological remarks

Trophic relationship between large scavengers like the cathartids here reported and the megafaunal mammal of South America have been Fig. 6. A) Anterior, B) medial, C) lateraland D) posterior views of tibiotarsus previously proposed (Tonni and Noriega, 1998; Alvarenga and Olson, CB-109. Scale bar equals 1 cm. 2004; Alvarenga et al., 2008; Jones et al., 2013, 2015; Mayr, 2017). Consequently, the causal of some scavenger species could be related to the late Pleistocene-Holocene megafaunal (Steadman and Martin, 1984; Hertel, 1995; Alvarenga and Olson, 2004;

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Fig. 7. Comparative anterior views of Cathartidae and Teratornithidae tibiotarsii. A) Teratornis merriami; B) CB-109; C) Vultur gryphus; D) Gymnogyps californianus;; E) E) Sarcoramphus papa. Abbreviations: canalis exten. = canalis extensorius, cond. lateralis = condylus lateralis, cond. medialis = condylus medialis, fov. lig. collat. lateralis = fovea ligamenti collateralis lateralis, incis. intercond. = incisura intercondylaris, tuber. distalis retin. extens. = tuberculum distalis retinaculi extensoris, tuber. proximalis retin. extens. = tuberculum proximalis retinaculi extensoris. Scale bar equals 1 cm.

Table 3 Tibiotarsus end width (in mm) in extant and fossil species (†).

Cathartidae sp. indet. CRB-109 Vultur gryphus Gymnogyps californianusa Sarcoramphus papa Pleistovultur nevesi† Gymnogyps howardae† Geronogyps reliquus† N = 3 N = 72 N = 4 N = 2

24 25.1–28.1 22.3–26.2 17.6–18.7 20.5 24.2–25.3 24.1

a -measurements taken from Fisher (1947).

Alvarenga et al., 2008; Jones et al., 2013; Jones et al., 2015; Galetti Author statement et al., 2018 and references therein). The access to large amounts of mammal carcasses promotes a great diversifications of feeding niches Washington Jones: Conceptualization, Investigation, Methodology, and evolutionary diversification of Cathartidae, which apparently Resources, Writing-Original Draft, Visualization. Andres´ Rinderknecht: reached its maximum during the Late Pleistocene on the American Conceptualization, Fossil Curation, Resources, Writing-Review & Edit­ continent (Emslie, 1987, 1988a, b; Tonni and Noriega, 1998). The ing, Visualization. Raúl Vezzosi: Resources, Investigation, Writing- associated megafauna and age context of the fossil cathartids reported Review & Editing. Felipe Montenegro: Resources, Writing-Review & here suggest the existence of this trophic relationship. Additionally, Editing. Martín Ubilla: Methodology, Resources, Writing-Review & Fisher (1946) identified a reversed direction of the process and the Editing. smoothness rami of the cathartid furcula as an adaptation to support and protect the relatively large and thin crop, not observed in other avian scavengers and birds of prey. Declaration of competing interest Currently, three size groups of scavenger birds are distributed in South America like genus Cathartes and Coragyps atratus (up to 2 kg, The authors declare that they have no known competing financial Dunning, 2008), the locally common medium size Sarcoramphus papa interests or personal relationships that could have appeared to influence (mean body mass 3.4 kg, Dunning, 2008), and the very large Andean the work reported in this paper. Condor Vultur gryphus (body mass range 9.6–13.6 kg, Dunning, 2008), nowadays confinedto the Andean and extra-Andean Patagonian region Acknowledgments (Lambertucci et al., 2008). Clearly, there exists a large gap of sizes be­ tween S. papa and V. gryphus. In this sense, the fossil cathartids reported We thank to the curators of the following institutional collections for – here could represent a group fillingthe gap between them (Tables 1 3). facilitating the use of the specimens studied and the use of photos and This implies a body size range not existent in living South American information under their care: Allison Farrell, Carrie M. Howard and scavenger bird guilds (Wallace and Temple, 1987; for an analysis of Kimball from (LACM); Christopher Milenski (USNM); Herculano North American avian scavengers see Hertel, 1995). This condition in­ Alvarenga (MHNT); Yolanda Davis and Federico Agnolin (MACN); Ser­ dicates a different combination of competitive interactions in the guild gio Bogan (CFA); Santiago Claramunt and Kevin Seymour (ROM). We of American avian scavengers during the Late Pleistocene (previously also thank to Marcos Cenizo for fossil cathartid and teratornithid pho­ proposed by Jones et al., 2013). tographs and literature. Specially thank to Rolando Bianchi (CB collec­ The fossil specimens reported here add new information to the re­ tion) for the access of the fossil material under his care and A. Rojas for cord of large New World vultures during the Late Pleistocene and pro­ the material in FC-DPV collection. The authors thank to Federico motes its prospection looking for new discoveries in Uruguayan Agnolin and two anonymous reviewers for your comments that clearly outcrops. The knowledge of the taxonomical diversity of avian scaven­ improved the manuscript. This work was funded by Programa de gers during the Quaternary seems to have just begun. Desarrollo de Ciencias Basicas´ (PEDECIBA), Agencia Nacional de Investigacion´ e Innovacion´ (ANII) and CSIC-Udelar (MU) of Uruguay.

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