Swiss Journal of Palaeontology

, Volume 138, Issue 2, pp 237–248 | Cite as

A new vertebrate continental assemblage from the Tortonian of Venezuela

  • Jorge D. Carrillo-BriceñoEmail author
  • Andrés E. Reyes-Cespedes
  • Rodolfo Salas-Gismondi
  • Rodolfo Sánchez
Regular Research Article


A wide variety of aquatic vertebrates from fluvio-lacustrine facies of northern South America (Colombia and Venezuela) have been used as unequivocal evidence to support hydrographic connections between western Amazonia and the Proto-Caribbean Sea during the Miocene. By the end of the Miocene, changes in the major hydrographic systems of the region produced losses of habitats and a regional faunal turnover, as has been documented in the geological record of the Urumaco region. Here, we report a new Tortonian aquatic and terrestrial vertebrate assemblage from two localities of the Caujarao Formation (El Muaco Member) in western Venezuela. The vertebrate assemblage includes a gharial (cf. †Gryposuchus pachakamue), alligatorid crocodylians (†Purussaurus and Alligatoridae indet.), a freshwater turtle (Chelus sp.), snakes (cf. Eunectes sp.), serrasalmids and pimelodids and thorny catfishes, a rodent (†Potamarchus sp.), pampatheres (†Scirrotherium sp.), sloths, as well as plant remains (coal and amber). Although the Caujarao Formation has been referred to as a fully marine environment, the new assemblage reported here suggests a freshwater input to the coastal area. Taxonomic and biogeographic affinities between the Muaco Member community and that reported from the Miocene proto-Amazonian systems are indicative of the persistence of ecological and hydrographic continuity at minimum until the end of the Miocene in at least an area of northwestern South America.


Neogene Miocene Caujarao Formation Orinoco River Biogeography 



We thank to Marcelo Sánchez-Villagra for academic support and for the financial assistance (University of Zurich research monies) that made this work possible. We especially thank the Taratara community, the Museo Ángel Segundo López and the Instituto del Patrimonio Cultural de Venezuela (IPC) for the authorization to collect and study the specimens. To the Alcaldía Bolivariana de Urumaco, the Universidad Experimental Francisco de Miranda, the Museo de Ciencias Naturales de Caracas, in Venezuela, the Mapuka Museum of Universidad del Norte, Colombia; the Natural History Museum of National University of San Marcos, Peru; and the Palaeontological Institute and Museum at the University of Zurich, Switzerland, for their valuable assistance and for access to comparative material. We greatly appreciate comments, suggestions and collaboration provided by Dr. Torsten Scheyer and Dr. Gabriel Aguirre-Fernández. The Editor and reviewers (Dr. Edwin Cadena, Dr. Leonardo Kerber, and Jaime Villafaña) are thanked for their valuable comments and suggestions on the manuscript.


This study was funded/support for the financial assistance of the University of Zurich research monies.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.


  1. Aguilera, O. A. (2010). Peces fósiles del Caribe de Venezuela. Washington: Gorham Printing.Google Scholar
  2. Aguilera, O. A., & Lundberg, J. G. (2010). Venezuelan Caribbean and Orinocoan Neogene fish. In M. R. Sánchez-Villagra, O. A. Aguilera, & F. Carlini (Eds.), Urumaco and Venezuelan Paleontology (pp. 129–152). Bloomington: Indiana Press University.Google Scholar
  3. Aguilera, O., Lundberg, J., Birindelli, J., Sabaj Pérez, M., Jaramillo, C., & Sánchez-Villagra, M. R. (2013). Palaeontological evidence for the last temporal occurrence of the ancient western Amazonian River outflow into the Caribbean. PLoS One,8, e76202. Scholar
  4. 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.Google Scholar
  5. Aguilera, O. A., Schwarzhans, W., & Béarez, P. (2016). Otoliths of the Sciaenidae from the Neogene of tropical America. Palaeo Ichthyologica,14, 7–90.Google Scholar
  6. Aguirre-Fernández, G., Carrillo-Briceño, J. D., Sánchez, R., Amson, E., & Sánchez-Villagra, M. R. (2017). fossil cetaceans (mammalia, cetacea) from the neogene of Colombia and Venezuela. Journal of Mammalian Evolution,24(1), 71–90.Google Scholar
  7. Albert, J. S., Lovejoy, N. R., & Crampton, W. G. R. (2006). Miocene tectonism and the separation of cis- and trans-Andean river basins: Evidence from Neotropical fishes. Journal of South American Earth Sciences,21(1), 14–27.Google Scholar
  8. Ameghino, F. (1891). Caracteres diagnósticos de cincuenta especies nuevas de mamíferos fósiles argentinos. Revista Argentina de Historia Natural,1, 129–167.Google Scholar
  9. Amson, E., Argot, C., McDonald, H. G., & de Muizon, C. (2015). Osteology and Functional Morphology of the Axial Postcranium of the Marine Sloth Thalassocnus (Mammalia, Tardigrada) with Paleobiological Implications. Journal of Mammalian Evolution,22(4), 473–518.Google Scholar
  10. Barbosa-Rodrigues, J. (1892). Les reptiles fossils de la Vallée de l’Amazone. Vellosia, contribuições do Museu Botânico do Amazonas,2, 41–60.Google Scholar
  11. Burmeister, G. H. (1885). Examen crítico de los mamíferos y reptiles fósiles denominados por D. Augusto Bravard y mencionados en su obra precedente. Annales del Museo Nacional de Buenos Aires,3, 93–174.Google Scholar
  12. Cadena, E., & Jaramillo, C. (2015a). Early to middle miocene turtles from the northernmost tip of South America: Giant testudinids, chelids, and podocnemidids from the Castilletes Formation, Colombia. Ameghiniana,52(2), 188–203.Google Scholar
  13. Cadena, E. A., & Jaramillo, C. (2015b). The first fossil skull of Chelus (Pleurodira: Chelidae, Matamata turtle) from the early Miocene of Colombia. Palaeontologia Electronica,18(2.32A), 1–10. Scholar
  14. Carrillo, J. D., Amson, E., Jaramillo, C., Sánchez, R., Quiroz, L., Cuartas, C., et al. (2018). The Neogene record of Northern South American native ungulates. Smithsonian Contributions to Palaeobiology,101, 1–67.Google Scholar
  15. Carrillo-Briceño, J. D., Carrillo, J. D., Aguilera, O. A., & Sanchez-Villagra, M. R. (2018). Shark and ray diversity in the Tropical America (Neotropics)—An examination of environmental and historical factors affecting diversity. PeerJ,6, e5313. Scholar
  16. Carrillo-Briceño, J. D., Maxwell, E., Aguilera, O. A., Sánchez, R., & Sánchez-Villagra, M. R. (2015). Sawfishes and other elasmobranch assemblages from the Mio-Pliocene of the South Caribbean (Urumaco Sequence, Northwestern Venezuela). PLoS One,10, e0139230. Scholar
  17. Dahdul, W. M. (2004). Fossil serrasalmine fishes (Teleostei, Characiformes) from the Lower Miocene of Northwestern Venezuela. Fossils of the Miocene Castillo Formation, Venezuela, Contributions on Neotropical Palaeontology, Special Papers in Palaeontology,71, 23–28.Google Scholar
  18. Delfino, M., & Sánchez-Villagra, M. (2018). A Late Miocene Pipine Frog from the Urumaco Formation, Venezuela. Ameghiniana,55(2), 210–214.Google Scholar
  19. Dentzien-Dias, P., Carrillo-Briceño, J. D., Francischini, H., & Sánchez, R. (2018). Paleoecological and taphonomical aspects of the Late Miocene vertebrate coprolites (Urumaco Formation) of Venezuela. Palaeogeography, Palaeoclimatology, Palaeoecology,490, 590–603.Google Scholar
  20. Díaz de Gamero, M. L. (1996). The changing course of the Orinoco River during the Neogene: A review. Palaeogeography, Palaeoclimatology, Palaeoecology,123(1), 385–402.Google Scholar
  21. Duméril, A. M. C. (1806). Zoologie analytique, ou méthode naturelle de classification des animaux, rendue plus facile a l’aide de tableux synoptiques. Paris: Allais.Google Scholar
  22. Edmund, G., & Theodor, J. (1997). A new giant Pampatheriid armadillo. In R. F. Kay, R. H. Madden, R. L. Cifelli, & J. J. Flynn (Eds.), Vertebrate paleontology in the neotropics, the Miocene fauna of La Venta, Colombia (pp. 227–232). Washington and London: Smithsonian Institution Press.Google Scholar
  23. Ferreira, G. S., Rincón, A. D., Solórzano, A., & Langer, M. C. (2016). Review of the fossil matamata turtles: Earliest well-dated record and hypotheses on the origin of their present geographical distribution. The Science of Nature,103(3), 28. Scholar
  24. Góis, F., Scillato-Yané, G. J., Carlini, A. A., & Guilherme, E. (2013). A new species of Scirrotherium Edmund & Theodor, 1997 (Xenarthra, Cingulata, Pampatheriidae) from the late Miocene of South America. Alcheringa: An Australasian Journal of Palaeontology,37(2), 177–188.Google Scholar
  25. González de Juana, C., Iturralde de Arozena, J. M., & Picard, C. X. (1980). Geología de Venezuela y de sus Cuencas Petrolíferas. Caracas: Ediciones Foninves.Google Scholar
  26. Hoorn, C., Wesselingh, F. P., ter Steege, H., Bermudez, M. A., Mora, A., Sevink, J., et al. (2010). Amazonia through time: Andean uplift, climate change, landscape evolution, and biodiversity. Science,330(6006), 927–931.Google Scholar
  27. Horovitz, I., Sanchez-Villagra, M. R., Vucetich, M. G., & Aguilera, O. A. (2010). Fossil rodents from the Late Miocene Urumaco and middle Miocene Cumaca formations, Venezuela. In M. R. Sánchez-Villagra, O. A. Aguilera, & F. Carlini (Eds.), Urumaco and Venezuelan paleontology (pp. 214–232). Bloomington: Indiana Press University.Google Scholar
  28. Hsiou, A. S., & Albino, A. M. (2009). Presence of the Genus Eunectes (Serpentes, Boidae) in the Neogene of Southwestern Amazonia, Brazil. Journal of Herpetology,43(4), 612–619.Google Scholar
  29. Hsiou, A. S., & Albino, A. M. (2010). New snake remains from the Miocene of northern South America. The Herpetological Journal,20(4), 249–259.Google Scholar
  30. Jaramillo, C. A., Hoorn, C., Silva, S. A. F., Leite, F., Herrera, F., Quiroz, L., et al. (2010). The origin of the modern Amazon rainforest: Implications of the palynological and palaeobotanical record. In C. Hoorn & F. P. Wesselingh (Eds.), Amazonia: Landscape and species evolution (pp. 334–337). Oxford: Blackwell Publishing.Google Scholar
  31. Kavanagh de Petzall, C. (1959). Estudio de una sección de la Formación Caujarao en el anticlinal de La Vela, Estado Falcón. Asociación Venezolana de Geólogos Mineros y Petroleros, Boletín Informativo,2, 269–319.Google Scholar
  32. Kerber, L., Negri, F. R., Ribeiro, A. M., Vucetich, M. G., & Souza-Filho, J. P. D. (2016). Late Miocene potamarchine rodents from southwestern Amazonia, Brazil—With description of new taxa. Acta Palaeontologica Polonica,61(1), 191–203.Google Scholar
  33. Langston, W. J. (1965). Fossil crocodilians from Colombia and the Cenozoic history of the Crocodilia in South America. University of California Publications in Geological Sciences,2, 1–157.Google Scholar
  34. Laurito, C. A., & Valerio, A. L. (2013). Scirrotherium antelucanus, una nueva especie de Pampatheriidae (Mammalia, Xenarthra, Cingulata) del Mioceno Superior de Costa Rica, América Central. Revista de Geología de América Central,49, 45–62.Google Scholar
  35. Lundberg, J. G. (1998). The temporal context for the diversification of Neotropical fishes. In L. Malabarba, R. E. Reis, R. P. Vari, C. A. S. de Lucena, & Z. M. S. de Lucena (Eds.), Phylogeny and Classification of Neotropical Fishes (pp. 49–68). Porto Alegre: Museu de Ciências e Tecnologia.Google Scholar
  36. Lundberg, J. G., Sabaj Pérez, M. H., Dahdul, W. M., & Aguilera, O. A. (2010). The Amazonian Neogene fish fauna. In C. Hoorn & F. P. Wesselingh (Eds.), Amazonia: Landscape and species evolution (pp. 281–301). Oxford: Blackwell Publishing.Google Scholar
  37. Moreno, F., Hendy, A. J. W., Quiroz, L., Hoyos, N., Jones, D. S., Zapata, V., et al. (2015). Revised stratigraphy of Neogene strata in the Cocinetas Basin, La Guajira, Colombia. Swiss Journal of Palaeontology,134(1), 5–43. Scholar
  38. 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. Scholar
  39. Quiroz, L., & Jaramillo, C. (2010). Stratigraphy and sedimentary environments of Miocene shallow to marginal marine deposits in the Urumaco Trough, Falcon Basin, western Venezuela. In M. R. Sánchez-Villagra, O. A. Aguilera, & F. Carlini (Eds.), Urumaco and Venezuelan Paleontology (pp. 153–172). Bloomington: Indiana Press University.Google Scholar
  40. Riff, D., & Aguilera, O. A. (2008). The world’s largest gharials Gryposuchus: Description of G. croizati n. sp. (Crocodylia, Gavialidae) from the Upper Miocene Urumaco Formation, Venezuela. Paläontologische Zeitschrift,82(2), 178–195.Google Scholar
  41. Riff, D., Romano, R. P. S., Oliveira, G. R., & Aguilera, O. A. (2010). Neogene Crocodile and Turtle Fauna in Northern South America. In C. Hoorn & F. P. Wesselingh (Eds.), Amazonia: Landscape and species evolution (pp. 259–280). Oxford: Blackwell Publishing.Google Scholar
  42. Rincón, A. D., Solórzano, A., Benammi, M., Vignaud, P., & McDonald, H. G. (2014). Chronology and geology of an Early Miocene mammalian assemblage in North of South America, from Cerro La Cruz (Castillo Formation), Lara State, Venezuela: Implications in the ‘changing course of Orinoco River’ hypothesis. Andean Geology,41(3), 507–528.Google Scholar
  43. Salas-Gismondi, R., Flynn, J. J., Baby, P., Tejada-Lara, J. V., Claude, J., & Antoine, P.-O. (2016). A new 13 million year old gavialoid crocodylian from proto-Amazonian mega-wetlands reveals parallel evolutionary trends in skull shape linked to longirostry. PLoS One,11, e0152453. Scholar
  44. Salas-Gismondi, R., Moreno-Bernal, J. W., Scheyer, T. M., Sánchez-Villagra, M. R., & Jaramillo, C. (2018). New Miocene Caribbean gavialoids and patterns of longirostry in crocodylians. Journal of Systematic Palaeontology. Scholar
  45. Sánchez-Villagra, M. R., Aguilera, O. A., & Carlini, F. (2010). Urumaco and Venezuelan Paleontology. Bloomington: Indiana Press University.Google Scholar
  46. Scheyer, T. M., Aguilera, O. A., Delfino, M., Fortier, D. C., Carlini, A. A., Sánchez, R., et al. (2013). Crocodylian diversity peak and extinction in the late Cenozoic of the northern Neotropics. Nature communications,4, 1907.Google Scholar
  47. Scheyer, T. M., & Delfino, M. (2016). The late Miocene caimanine fauna (Crocodylia: Alligatoroidea) of the Urumaco Formation, Venezuela. Palaeontologia Electronica,19(3.48A), 1–57. Scholar
  48. Smith, C. J., Collins, L. S., Jaramillo, C., & Quiroz, L. (2010). Marine paleoenvironment of Miocene–Pliocene formations of north-central Falcón state, Venezuela. Journal of Foraminiferal Research,40(3), 266–282.Google Scholar
  49. Vallenilla, L. P. (1961). Estratigrafía de las formaciones Caujarao, La Vela y Coro en sus localidades tipo, Estado Falcón. Asociación Venezolana de Geología, Minería y Petroleo, Boletín Informativo,4(2), 29–79.Google Scholar
  50. Wagler, J. G. (1830). Natürliches System der Amphibien: mit vorangehender Classification der Säugethiere und Vögel: ein Beitrag zur vergleichenden Zoologie. München In der J.G. Cotta’scchen Buchhandlung.Google Scholar
  51. Wood, R. C. (1976). Two new species of Chelus (Testudines: Pleurodira) from the Late Tertiary of northern South America. Breviora,435, 1–26.Google Scholar
  52. Wozniak, J., & Wozniak, M. H. (1987). Bioestratigrafía de la región nor-central de la Serranía de Falcón, Venezuela nor-occidental. Boletín de Geología (Caracas, Venezuela),16, 101–139.Google Scholar

Copyright information

© Akademie der Naturwissenschaften Schweiz (SCNAT) 2018

Authors and Affiliations

  1. 1.Universität Zürich, Paläontologisches Institut und MuseumZurichSwitzerland
  2. 2.Departamento de Física y GeocienciasUniversidad del NorteBarranquillaColombia
  3. 3.BioGeoCiencias Lab, Facultad de Ciencias y Filosofía/CIDISUniversidad Peruana Cayetano HerediaLimaPeru
  4. 4.Departamento de Paleontología de VertebradosMuseo de Historia Natural, UNMSMLimaPeru
  5. 5.Museo Paleontológico de UrumacoUrumacoVenezuela

Personalised recommendations