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The Crocodylomorph Fauna of the Cenozoic of South America and Its T Evolutionary History: a Review ∗ Giovanne M

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The Crocodylomorph Fauna of the Cenozoic of South America and Its T Evolutionary History: a Review ∗ Giovanne M Journal of South American Earth Sciences 90 (2019) 392–411 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames The crocodylomorph fauna of the Cenozoic of South America and its T evolutionary history: a review ∗ Giovanne M. Cidadea, , Daniel Fortierb, Annie Schmaltz Hsioua a Laboratório de Paleontologia, Departamento de Biologia, FFCLRP, Universidade de São Paulo, Avenue Bandeirantes, 3900, Ribeirão Preto, São Paulo, 14040-901, Brazil b Campus Doutor Amílcar Ferreira Sobral, Universidade Federal do Piauí, Floriano, Piauí, Brazil ARTICLE INFO ABSTRACT Keywords: The fossil crocodylomorph fauna of the Cenozoic of South America is very rich and diverse. Historically, few Crocodylomorpha publications have been dedicated to providing an overall review of this fauna, with most reviews focusing on South America specific areas. However, the fact that many new species, taxonomic reviews and description of new specimens Cenozoic have been proposed in the last decade makes a comprehensive review of the fossil crocodylomorph fauna of the Sebecidae South American Cenozoic necessary. The only crocodylomorph lineages to have a fossil record comprising Late Caimaninae Cretaceous and Cenozoic deposits in South America is the Dyrosauridae. Sebecidae or its predecessors, however, Dyrosauridae are very likely to have inhabited the continent during the Cretaceous-Paleogene transition as well; both Dyrosauridae and Sebecidae are considered here to have survived the Cretaceous-Paleogene extinction while inhabiting South America. Caimaninae (Alligatoroidea) arrived either in the late Cretaceous or in the early Paleocene coming from North America. The fossil record of Caimaninae is present, however, only from the Paleocene. By the Eocene, there are no records of Dyrosauridae in South America; this group was globally extinct after the Eocene, possibly due to the global cooling that occurred by the end of the epoch. Sebecids and cai- manines solely comprised the crocodylomorph fauna of the continent until the Miocene, where there are the first records of Gavialoidea (Gryposuchinae) and a possible first dispersion of Crocodyloidea, through the tentative tomistomines Charactosuchus and Brasilosuchus. Gryposuchinae likely arrived in the continent from Africa or from Asia during the Oligocene. Charactosuchus and Brasilosuchus may have come from North America where tomistomines lived from the Oligocene to the Pliocene. Sebecids were extinct after the middle Miocene; Charactosuchus, Gryposuchinae, Purussaurus, Mourasuchus and durophagous caimanines such as Gnatusuchus were extinct after the late Miocene. These extinctions are related to changes in the drainage basins caused by elevation of the Andes mountain range. Only the extant caimanine Caiman, Melanosuchus and Paleosuchus would survive the Miocene, being enriched by Crocodylus from the Pliocene onwards, which is likely that Crocodylus arrived in the American continents from Africa. The current fossil record indicates that these four genera comprise the extant crocodylian fauna of the continent since the Pliocene. Although our knowledge on croco- dylomorph fossil fauna of the South American Cenozoic has increased continuosly, especially in the last decade, much yet must be done, especially on the taxonomy and phylogeny of Brasilosuchus and Charactosuchus, field- works in Paleogene and Pliocene localities, and the evolution of the specialized Caimaninae morphotypes. Abbreviations: American Musem of Natural History, New York, United States, AMNH; Colección de Paleontología de Vertebrados de la Alcaldía de Urumaco, Urumaco, Venezuela, AMU CURS; Centro de Investigaciones Antropológicas, Arqueológicas y Paleontológicas, Universidad Nacional Experimental Francisco de Miranda, Coro, Venezuela, CIAAP-UNEFM; Departamento de Geologia, Centro de Tecnologia e Geociências, Universidade Federal de Pernambuco, Recife, Brazil, DG- CTG-UFPE; Museo Geológico, Instituto Nacional de Investigaciones en Geociencias, Minería y Quimica, Bogotá, Colombia, IGM; Museu de Ciências da Terra, Serviço Geológico Nacional, Rio de Janeiro, Brazil, MCT; Museo Argentino de Ciéncias Naturales, Buenos Aires, Argentina, MACN; Museo de Ciéncias Naturales de Caracas, Caracas, Venezuela, MCNC; Museum of Comparatize Zoology, Cambridge, United States, MCZ; Museo de La Plata, La Plata, Argentina, MLP; Museo Paleontológico Egidio Feruglio, Trelew, Argentina, MPEF; Museo de Historia Natural de San Marcos, Lima, Peru, MUSM; Instituto Miguel Lillo, Tucumán, Argentina, PVL; University of California Museum of Paleontology, Berkeley, California, UCMP; Florida Musem of Natural History, University of Florida, Gainesville, United States, UF; Universidade Federal do Acre, Rio Branco, Brazil, UFAC; Universidade Federal Uberlândia, campus Monte Carmelo, Brazil, UFU ∗ Corresponding author. E-mail addresses: [email protected] (G.M. Cidade), [email protected] (D. Fortier), [email protected] (A.S. Hsiou). https://doi.org/10.1016/j.jsames.2018.12.026 Received 4 September 2018; Received in revised form 26 December 2018; Accepted 26 December 2018 Available online 27 December 2018 0895-9811/ © 2018 Elsevier Ltd. All rights reserved. G.M. Cidade et al. Journal of South American Earth Sciences 90 (2019) 392–411 1. Introduction Although previously published works have offered comprehensive revisions of such record (e.g. Gasparini, 1996; Figueiredo and Pinheiro, Crocodylomorphs have a very taxonomically rich and morphologi- 2011), many have focused either on specific areas or countries, such as cally diverse fossil record throughout the Cenozoic of South America. the Amazon region (Riff et al., 2010; Scheyer and Moreno-Bernal, These include not only fossils assigned to the crocodylian genera that 2010), Argentina (Gasparini, 1981; Bona et al., 2013a; Bona and currently inhabit the continent (Patterson, 1936; Medina, 1976; Souza- Barrios, 2015) and Brazil (Riff et al., 2012), and, as a result, compre- Filho, 1987; Salas-Gismondi et al., 2007, 2015; Fortier et al., 2009, hensive revisions of the crocodylomorph fossil fauna of the Cenozoic of 2014; Bona and Carabajal, 2013; Bona et al., 2013a, 2014; 2017a; South America are scarce. Additionally, many new fossil species and Scheyer et al., 2013; Foth et al., 2017), but also a myriad of exclusively revisions of previously proposed species have been recently published fossil taxa (e.g. Rovereto, 1912; Rusconi, 1933, 1935; 1937; Patterson, (Bona et al., 2013a, 2013b; 2017a, 2017b; 2018; Pinheiro et al., 2013; 1943; Simpson, 1937a, 1937b; Langston, 1965; Langston and Gasparini, Scheyer et al., 2013; Fortier et al., 2014; Kellner et al., 2014; Hastings 1997; Aguilera, 2004; Paolillo and Linares, 2007; Riff et al., 2010, et al., 2015; Salas-Gismondi et al., 2015, 2016; 2018; Scheyer and 2012; Scheyer and Moreno-Bernal, 2010; Bona et al., 2013a; Bona and Delfino, 2016; Cidade et al., 2017, 2018a; Foth et al., 2017), justifying, Barrios, 2015; Salas-Gismondi et al., 2015; Scheyer and Delfino, 2016), thus, the need for this comprehensive revision. Furthermore, an over- including some that possess ecomorphotypes that are absent in the view of the biogeography, faunal successions and paleoecology of the current continental crocodylian fauna. different groups that comprise the crocodylomorph fossil fauna ofthe These morphotypes include the terrestrial, top predators sebecids Cenozoic of South America is also be provided. (Simpson, 1937a; Colbert, 1946; Langston, 1965; Gasparini, 1972; Buffetaut and Hoffstetter, 1977; Paolillo and Linares, 2007; Molnar, 2. Results 2010, 2013; Kellner et al., 2014; Molnar and Vasconcellos, 2016), the longirrostrine, marine, estuarine and freshwater inhabitants Dyrosaur- 2.1. Crocodylomorpha of the Paleogene idae (Cope, 1886; Barbosa et al., 2008; Andrade and Sayão, 2014; Hastings et al., 2010, 2011, 2015), the giant, semi-aquatic top predator When K-Pg extinction occurred, around 66 Ma ago (Renne et al., Purussaurus (Barbosa-Rodrigues, 1892; Nopcsa, 1924; Mook, 1941; 2013), only one out of seven crocodylomorph lineages to survive al- Langston, 1965; Bocquentin-Villanueva et al., 1989; Aguilera et al., ready inhabited South America: Dyrosauridae (Fig. 1), which already 2006; Aureliano et al., 2015), the giant, flat-headed “duck faced” pu- has a record in the Late Cretaceous of the continent (Langston, 1965; tative “gulp-feeder” Mourasuchus (Price, 1964; Langston, 1965, 1966; Hastings et al., 2015). The clade Sebecidae (Fig. 2) has a large like- 2008; Bocquentin-Villanueva, 1984; Gasparini, 1985; Bocquentin and lihood of being inhabiting South America during the K-Pg transition as Souza-Filho, 1990; Cidade et al., 2017), highly adapted durophagous well. Although Sebecidae itself is traditionally proposed as an ex- caimanines like Gnatusuchus, Kuttanacaiman and Globidentosuchus, clusively Cenozoic group (see Pol and Powell, 2011), it has been pro- among others (Langston, 1965; Salas-Gismondi et al., 2015) and the posed to be closely related either to Baurusuchidae, forming the clade longirrostrine, predominantely piscivorous gavialoids (Gryposuchinae, Sebecosuchia (e.g. Turner and Calvo, 2005; Pol and Powell, 2011) or which also included large-sized taxa such as Gryposuchus; Gürich, 1912; Peirosauridae, forming the clade Sebecia (e.g. Larsson and Sues, 2007). Langston, 1965; Gasparini, 1968; Sill, 1970; Bocquentin-Villanueva and However, as both of these clades are predominantly from South Buffetaut, 1981; Buffetaut,
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