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Type: Article
Published: 2019-08-14
Page range: 475–486
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Taxonomic review of two fossil crocodylians from the Cenozoic of South America and its implications for the crocodylian fauna of the continent

Laboratório de Paleontologia, Departamento de Biologia, FFCLRP, Universidade de São Paulo. Avenue Bandeirantes, 3900, Ribeirão Preto, São Paulo, Brazil, 14040-901. Telephone: +5516 3315 0446.
Laboratório de Geociências e Paleontologia, Núcleo de História Natural, CAFS, Universidade Federal do Piauí, BR 343, km 3,5, Floriano, Piauí, Brazil, 64808-605.
Laboratorio de Paleontología, Centro de Ecología, Instituto Venezolano de Investigaciones Científicas; Carretera Panamericana, Caracas, Venezuela.
Laboratório de Paleontologia, Departamento de Biologia, FFCLRP, Universidade de São Paulo. Avenue Bandeirantes, 3900, Ribeirão Preto, São Paulo, Brazil, 14040-901. Telephone: +5516 3315 0446.
Reptilia El Breal de Orocual-Honda Group-Miocene-Pliocene-Pleistocene-Caimaninae


The crocodylomorph fauna of the Cenozoic of South America is one of the richest and most diverse in the world. The most diverse group within that fauna is Alligatoroidea, with nearly all of its species belonging to the Caimaninae clade. Many of the fossil alligatoroid species from the Cenozoic of South America were proposed based on very incomplete remains, and as a result their validity requires revision. Two such species are Balanerodus logimus Langston, 1965, from the middle Miocene of Colombia and Peru, and Caiman venezuelensis Fortier & Rincón, 2012, from the Pliocene-Pleistocene of Venezuela. This study has performed a thorough review of the taxonomic status of these two alligatoroid species, concluding that B. logimus is a nomen dubium and that Ca. venezuelensis is a junior synonym of the extant species Ca. crocodilus. This review offers a significantly more accurate scenario for alligatoroid diversity in the Cenozoic of South America in different epochs such as the Miocene and Pleistocene. Additionally, the record of Ca. crocodilus from the Pleistocene of Venezuela is the first fossil record that can be assigned to this species.



  1. Agassiz, L. (1849) Remarks on fossil crocodiles. Proceedings of the Academy of Natural Sciences of Philadelphia, 4 (1848–1849), 169.

    Aguilera, O.A., Riff, D. & Bocquentin-Villanueva, J. (2006) A new giant Purussaurus (Crocodyliformes, Alligatoridae) from the upper Miocene Urumaco Formation, Venezuela. Journal of Systematic Palaeontology, 4 (3), 221–232.

    Aureliano, T., Ghilardi, A.M., Guilherme, E., Souza-Filho, J.P., Cavalcanti, M. & Riff, D. (2015) Morphometry, bite-force, and paleobiology of the Late Miocene Caiman Purussaurus brasiliensis. Plos One, 10 (2), e0117944.

    Ayarzagüena, J. (1984) Ecología del caimán de anteojos (Caiman crocodilus L.) en los Llanos de Apure (Venezuela). Doñana Acta Vertebrata, 10 (3), 1–136.

    Benton, M.J. & Clark, J.M. (1988) Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M.J. (Ed.), The Phylogeny and Classification of the Tetrapods. Clarendon Press, Oxford, pp. 295–338.

    Barbosa-Rodrigues, B. (1892) Les Reptiles fossiles de la vallée de l’Amazone. Vellosia, 2, 41–46.

    Bona, P., Riff, D. & Gasparini, Z. (2013) Late Miocene crocodylians from Northeast Argentina: new approaches about the austral components of the Neogene South American crocodylian fauna. Transactions of the Royal Society of Edinburgh. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 103, 1–20.

    Bona, P., Ezcurra, M.D., Barrios, F. & Fernandez-Blanco, M.V. (2018) A new Palaeocene crocodylian from southern Argentina sheds light on the early history of caimanines. Proceedings of the Royal Society B, 285, 20180843.

    Brochu, C.A. (1997) Phylogenetic Systematics and Taxonomy of Crocodylia., Unpublished PhD Thesis, University of Texas, Austin, 432 pp.

    Brochu, C.A. (1999) Phylogenetics, taxonomy, and historical biogeography of Alligatoroidea. Memoir of the Society of Vertebrate Paleontology, 6, 9–100.

    Brochu, C.A. (2003) Phylogenetic approaches toward crocodilian history. Annual Review of Earth and Planetary Sciences, 31, 357–397.

    Brochu, C.A. (2004) Alligatorine phylogeny and the status of Allognathosuchus Mook, 1921. Journal of Vertebrate Paleontology, 24, 856–872.

    Brochu, C.A. (2010) A new alligatoroid from the Lower Eocene Green River Formation of Wyoming and the origin of caimans. Journal of Vertebrate Paleontology, 30, 1109–1126.

    Brochu, C.A. (2011) Phylogenetic relationships of Necrosuchus ionensis Simpson, 1937 and the early history of caimanines. Zoological Journal of the Linnean Society, 163, S228–S256.

    Case, E.C. (1925) Note on a new species of the Eocene crocodilian Allognathosuchus, A. wartheni. Contributions from the Museum of Geology, University of Michigan, 2, 93–97.

    Cossette, A.P. & Brochu, C.A. (2018) A new specimen of the alligatoroid Bottosaurus harlani and the early history of character evolution in alligatorids, Journal of Vertebrate Paleontology, 38 (4), 1–22.

    Cidade, G.M. & Hsiou, A.S. (2018) New morphological, evolutionary and paleoecological interpretations on the genus Eocaiman (Crocodylia, Caimaninae) from the Cenozoic of South America. In: Marzola, M., Mateus, O. & Moreno-Azanza, M. (Eds.), Abstract book of the XVI Annual Meeting of the European Association of Vertebrate Palaeontology. Departamento de Ciências da Terra, Faculdade Ciências e Tecnologia, Universidade Nova de Lisboa, Caparica, p. 46.

    Cidade, G.M., Solórzano, A., Rincón, A.D., Riff, D. & Hsiou, A.S. (2017) A new Mourasuchus (Alligatoroidea, Caimaninae) from the late Miocene of Venezuela, the phylogeny of Caimaninae and considerations on the feeding habits of Mourasuchus. PeerJ, 5, e3056.

    Cidade, G.M., Fortier, D. & Hsiou, A.S. (2019) The crocodylomorph fossil record of the Cenozoic of South America and its evolutionary history: a review. Journal of South American Earth Sciences, 90, 392–411.

    Cuvier, G.L. (1807) Sur les differentes especes de crocodiles vivan set sur leur caracteres distinctifs. Annales du Museum d’Histoire Naturelle, 10, 8–66.

    Daudin, F.M. (1802) Histoire naturelle, generale et particuliere, des reptiles. Paris: Del’imprimerie de F. Dufort, Volume 2, Paris, 432 pp.

    Erickson, B.R. (1972) Albertochampsa langstoni, gen. et sp. nov., a new alligator from the Cretaceous of Alberta. Scientific Publications of the Science Museum of Minnesota, New Series, 2, 1–13.

    Escobedo-Galván, A.H., Cupul-Magaña, F.G. & Velasco, J.A. (2011) Misconceptions about the taxonomy and distribution of Caiman crocodilus chiapasius and C. crocodilus fuscus (Reptilia: Crocodylia: Alligatoridae). Zootaxa, 3015, 66–68.

    Escobedo-Galván, A.H., Velasco, J.A., González-Maya, J.F. & Resetar, A. (2015) Morphometric analysis of the Rio Apaporis caiman (Reptilia, Crocodylia, Alligatoridae). Zootaxa, 4059 (3), 541–554.

    Fortier, D., Brochu, C.A. & Souza-Filho, J.P. (2009) The oldest record of Caiman yacare. Journal of Vertebrate Paleontology, 29 (3), 97A.

    Fortier, D. & Rincón, A. (2012) Pleistocene crocodylians from Venezuela, and the description of a new species of Caiman. Quaternary International, 305, 141–148.

    Fortier, D.C., Souza-Filho, J.P., Guilherme, E., Maciente, A. & Schultz, C.L. (2014) A new specimen of Caiman brevirostris (Crocodylia, Alligatoridae) from the Late Miocene of Brazil. Journal of Vertebrate Paleontology, 34 (4), 820–834.

    Gilmore, C.W. (1911) A new fossil alligator from the Hell Creek beds of Montana. Proceedings of the United States National Museum, 41, 297–302.

    Gilmore, C.W. (1946) A new crocodilian from the Eocene of Utah. Journal of Paleontology, 20 (1), 62–67.

    Gmelin, J. (1789) Linnei Systema Naturae. G. E. Beer, Leipzig, 1057 pp.

    Gorzula, S. (1994). A longirostrine Caiman crocodilus from central Venezuela. Crocodile Specialist Group Newsletter, 13 (3), 16.

    Gray, J.E. (1844) Catalogue of Tortoises, Crocodilians, and Amphisbaenians in the Collection of the British Museum. British Museum (Natural History), London, 88 pp.

    Kälin, J.A. (1940) Arambourgia nov. gen. gaudryi de Stefano sp., ein kurzschnauziger Crocodilide aus den Phosphoriten des Quercy. Eclogue Geologicae Helvetiae, 32, 185–186.

    Langston, W. (1965) Fossil crocodilians from Colombia and the Cenozoic history of the Crocodilia in South America. University of California Publications in Geological Sciences, 52, 1–168.

    Langston, W. & Gasparini, Z. (1997) Crocodilians, Gryposuchus, and the South Americans gavials. In: Kay, R.F., Madden, R.H., Ciffelli, R.L. & Flynn, J.J. (Eds.), Vertebrate paleontology in the neotropics: the Miocene fauna of La Venta, Colombia. Smithsonian Institution, Washington, pp. 113–154.

    Laurenti, J.N. (1768) Specimen Medicum, Exhibens Synopsin Reptilium Emendatum cum Experimentis Circa Venena et Antidota Reptilium Austriacorum. Joan. Thomae nob. de Trattnern, Vienna, 217 pp.

    Linnaeus, C. (1758) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Salvius, Stockholm, 824 pp.

    Martin, J.E. (2010) A new species of Diplocynodon (Crocodylia, Alligatoroidea) from the Late Eocene of the Massif Central, France, and the evolution of the genus in the climatic context of the Late Palaeogene. Geological Magazine, 147, 596–610.

    Martin, J.E. & Lauprasert, K. (2010) A new primitive alligatorine from the Eocene of Thailand: relevance of Asiatic members to the radiation of the group. Zoological Journal of the Linnean Society, 158, 608–628.

    Martin, J.E., Smith, T., Broin, F.L., Escuillié, F. & Delfino, M. (2014) Late Palaeocene eusuchian remains from Mont de Berru, France, and the origin of the alligatoroid Diplocynodon. Zoological Journal of the Linnean Society, 172, 867–891.

    Medem, F. (1955) A new subspecies of Caiman sclerops from Colombia. Fieldiana: Zoology, 37, 339–344.

    Meyer, H. von. (1832) Paleologica zur Geshichte der Erde und irher Geschöpfe. S. Schmerber, Frankurt-am-Main, 560 pp.

    Mook, C.C. (1921) Allognathosuchus, a new genus of Eocene crocodilians. Bulletin of the American Museum of Natural History, 44, 105–110.

    Mook, C.C. (1941) A new crocodilian, Hassiacosuchus kayi, from the Bridger Eocene Beds of Wyoming. Annals of the Carnegie Museum, 28, 207–220.

    Moreno-Bernal, J.W., Head, J. & Jaramillo, C.A. (2016) Fossil crocodilians from the high Guajira Peninsula of Colombia: Neogene faunal change in northernmost South America. Journal of Vertebrate Paleontology, 36 (3), e1110586.

    Norell, M.A. (1988) Cladistic approaches to evolution and paleobiology as applied to the phylogeny of alligatorids. Unpublished PhD Thesis, Yale University, New Haven, 272 pp.

    Norell, M.A., Clark, J.M. & Hutchison, J.H. (1994) The Late Cretaceous alligatoroid Brachychampsa montana (Crocodylia): new material and putative relationships. American Museum Novitates, 3116, 1–26.

    Oaks, J.R. (2011) A time-calibrated species tree of Crocodylia reveals a recent radiation of the true crocodiles. Evolution, 65, 3285–3297.

    Okamoto, K.W., Langerhans, R.B., Rashid, R. & Amarasekare, P. (2015) Microevolutionary patterns in the common caiman predict macroevolutionary trends across extant crocodilians. Biological Journal of the Linnean Society, 116, 834–846.

    Onary, S., Rincón, A.D. & Hsiou, A.S. (2018) Fossil snakes (Squamata, Serpentes) from the tar pits of Venezuela: taxonomic, palaeoenvironmental, and palaeobiogeographical implications for the North of South America during the Cenozoic/Quaternary boundary. PeerJ 6 e5402.

    Riff, D., Romano, P.S.R., Oliveira, G.R. & Aguilera, O.A. (2010) Neogene Crocodile and Turtle Fauna in Northern South America. In: Hoorn, C. & Wesselingh, F. (Eds.), Amazonia, Landscape and Species Evolution: A Look into the Past. Wiley-Blackwell, London, pp. 259–280.

    Rincón, A.D., Parra, G.E., Prevosti, F.J., Alberdi, M.T. & Bell, C.J. (2009) A preliminary assessment of the mammalian fauna from the Pliocene–Pleistocene El Breal de Orocual locality, Monaga State, Venezuela. Bulletin of the Museum of Northern Arizona, 65, 593–620.

    Rio, J.P., Mannion, P.D., Tschopp, E., Martin, J.E. & Delfino, M. (2019) Reappraisal of the morphology and phylogenetic relationships of the alligatoroid crocodylian Diplocynodon hantoniensis from the Late Eocene of the United Kingdom. Zoological Journal of the Linnean Society, zlz034.

    Salas-Gismondi, R., Antoine, P.O., Baby, P., Brusset, S., Benammi, M., Espurt, N., De Franceschi, D., Pujos, F., Tejada, J. & Urbina, M. (2007) Middle Miocene Crocodiles from the Fitzcarrald Arch, Amazonian Peru. Cuadernos del Museo Geominero, 8, 355–360.

    Salas-Gismondi, R., Flynn, J.J., Baby, P., Tejada-Lara, J.V., Wesselingh, F.P. & Antoine, P.O. (2015) A Miocene hyperdiverse crocodylian community reveals peculiar trophic dynamics in proto-Amazonian mega-wetlands. Proceedings of the Royal Society, 282, 20142490.

    Scheyer, T.M., Aguilera, O.A., Delfino, M., Fortier, D.C., Carlini, A.A., Sánchez, R., Carrillo-Briceño, J.D., Quiroz, L. & Sánchez-Villagra, M.R. (2013) Crocodylian diversity peak and extinction in the late Cenozoic of the northern Neotropics. Nature Communications, 4, 1907.

    Schneider, J.G. (1801) Historiae Amphibiorum naturalis et literariae. Fasciculus secundus continens Crocodilos, Scincos, Chamaesauras, Boas. Pseudoboas, Elapes, Angues. Amphisbaenas et Caecilias. Frommanni, Jena, 364 pp.

    Simpson, G.G. (1933) A new crocodile from the Notostylops beds of Patagonia. American Museum Novitates, 623, 1–9.

    Skutschas, P., Danilov, I.G., Kodrul, T.M. & Jin, J. (2014) The first discovery of an alligatorid (Crocodylia, Alligatoroidea, Alligatoridae) in the Eocene of China. Journal of Vertebrate Paleontology, 34, 471–476.

    Solórzano, A., Rincón, A.D., Cidade, C.M., Núñez-Flores, M. & Sánchez, L. (2019) Lower Miocene alligatoroids (Crocodylia) from the Castillo Formation, northwest of Venezuela. Palaeobiodiversity and Palaeoenvironments, 99, 241–259.

    Souza-Filho, J.P. (1987) Caiman brevirostris sp. nov., um novo Alligatoridae da Formação Solimões (Pleistoceno) do Estado do Acre, Brasil. Anais X Congresso Brasileiro de Paleontologia, Rio de Janeiro, 1, 173–180.

    Spix, J.B. (1825) Animalia nova sive species novae Lacertarum, quas in itinere per Brasiliam annis MDCCCXVII-MDCCCXX jussu et auspiciis Maximiliani Josephi I. Bavaria Regis suscepto collegit et descripsit D. J. B. De Pix. Typis Franc. Seraph. Hübschamanni, Munich, 26 pp.

    Stefano, G., (1905) Appunti sui Batraci e rettili del Quercy appartenenti alla collezioni Rossignol. Bolletino della Societa Geologia Italiana, 24, 17–67.

    Venegas-Anaya, M., Crawford, A.J., Escobedo-Galván, A.H., Sanjur, O.I., Densmore, L.D. & Bermingham, E. (2008) Mitochondrial DNA phylogeography of Caiman crocodilus in Mesoamerica and South America. Journal of Experimental Zoology, 309A, 614–627.

    Wang, Y.-Y., Sullivan, C. & Liu, J., (2016) Taxonomic revision of Eoalligator (Crocodylia, Brevirostres) and the paleogeographic origins of the Chinese alligatoroids. PeerJ, 4, e2356.

    Weitzel, K. (1935) Hassiacosuchus haupti n.g n.s. ein durophages Krokodil aus dem Mitteleozän von Messel. Notizblatt des Vereins für Erdkunde und der hessischen geologischen Landesanstalt zu Darmstadt, 16, 40–49.

    Wood, S.V. (1846) On the discovery of an alligator and of several new Mammalia in the Hordwell Cliff; with observations upon the geological phenomena of that locality. London Geological Journal, 1, 117–122.

    Wu, X.C., Brinkman, D.B. & Russell, A.P. (1996) A new alligator from the Upper Cretaceous of Canada and the relationships of early eusuchians. Palaeontology, 39 (2), 351–375.