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A new fossil canid assemblage from the late Pleistocene of northern South America: the canids of the Inciarte asphalt pit (Zulia, Venezuela), fossil record and biogeography

Published online by Cambridge University Press:  14 July 2015

Francisco J. Prevosti
Affiliation:
Departamento Cientifico Paleontologia Vertebrados, Museo de La Plata, Paseo del Bosque S/N°, 1900 La Plata, Buenos Aires, Argentina, Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET),
Ascanio D. Rincón
Affiliation:
Lab. Biología de Organismos, Centro de Ecología, Instituto Venezolano de Investigaciones Cientificas (IVIC), km 11 de la Carretera Panamericana, Apartado Postal 21827, Caracas 1020-A Venezuela,

Abstract

Canids invaded South American during the Pliocene (2.8–3.1 Ma BP) as part of the Great American Biotic Interchange, but their record on the continent is relatively poor until the Pleistocene. However, even the Pleistocene record of the group is patchy and biased, with few records for the northern part of South America. In this paper we describe a new assemblage of canids found at the Inciarte asphalt pits (Zulia, Venezuela). These remains are associated with a variety of extinct Pleistocene mammals dated between 25–27 Ka BP. Canids in this assemblage include Protocyon troglodytes (Lund 1838), Protocyon sp., Canis dirus Leidy, 1858, Urocyon cf. U. cinereoargenteus (Schreber, 1775), and an indeterminate canid. This is the first record of Urocyon Baird, 1758 from the Late Pleistocene in South America and constrains the timing of its dispersal to South America. The distributional record of Protocyon Giebel, 1855 and P. troglodytes is expanded 1,500 km north of previous records and constitutes the first record of these taxa in Venezuela. The presence of C. dirus at Inciarte constitutes the fourth locality for this species in South America.

Type
Research Article
Copyright
Copyright © The Paleontological Society 

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References

Allen, J. A. 1876. Description of some remains of an extinct species of wolf and extinct species of deer from the lead region of of the upper Mississippi. American Journal of Science, ser. 3, 11:4751.Google Scholar
Ameghino, F. 1882. Catálogo de las colecciones de antropología prehistórica y de paleontología de Florentino Ameghino, Partido de Mercedes. Catálogo de la Sección de la Provincia de Buenos Aires (República Argentina), en la Exposición Continental Sudamericana, Anexo A: 3542.Google Scholar
Ameghino, F. 1902. Notas sobre algunos mamíferos fósiles nuevos o poco conocidos del valle de Tarija. Anales del Museo Nacional de Buenos Aires, 3:225261.Google Scholar
Ameghino, F. 1906. La perforación astragaliana en Priodontes, Canis (Chrysocyon) y Typotherium . Anales del Museo de Buenos Aires, 6:119.Google Scholar
Arata, A. A. 1959. Revaluation of the Pleistocene Urocyon seminolensis from Florida. Quarterly Journal of Florida Academy of Science, 22:133.Google Scholar
Baird, S. F. 1757 [1758]. Mammals, p. 1757. In Nicholson, A. O. P. (ed.), Reports of explorations and surveys to ascertain the most practicable and economical route for a railroad from the Mississippi River to the Pacific Ocean. Senate Executive Document no. 78, Vol. 8. (Pt. 1), Washington DC.Google Scholar
Berman, W. D. 1994. Los carnívoros continentales (Mammalia, Carnivora) del Cenozoico en la provincia de Buenos Aires. Unpublished doctoral dissertation, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, 418 p.Google Scholar
Berta, A. 1987. Origin, diversification, and zoogeography of the South American Canidae. Fieldiana Zoology (n. s.), 39:455471.Google Scholar
Berta, A. 1988. Quaternary evolution and biogeography of the large South American Canidae (Mammalia: Carnivora). University of California Publication in Geological Sciences, 132:1149.Google Scholar
Bisbal, F. 1982. Nuevos registros de distribución para el zorro gris Urocyon cinereoargenteus venezuelae (Mammalia, Carnivora). Acta Científica Venezuelica, 33:255257.Google Scholar
Bocquentin, J. 1979. Mammifères fossiles du Pléistocène supérieur de Muaco, État de Falcón, Vénézuéla. Unpublished doctoral dissertation, L'Université Pierre et Marie Curie, Paris, 112 p.Google Scholar
Bowdich, T. E. 1821. An Analysis of the Natural Classifications of Mammalia for the Use of Students and Travelers. J. Smith, Paris, 115 p.Google Scholar
Buchalczyk, T., Dynowski, J., and Szteyn, S. 1981. Variations in number and asymmetry of the skull in the wolf. Acta Theriologica, 26:2330.Google Scholar
Burmeister, H. 1856. Erläuterungen sur Fauna Brasiliens, Enthaltend Abbildungen und Ausführliche Beschreibungen und Schreibungen Neuer Oder Ungenügend Bekannter Thier-Arten. George Reimer, Berlin, 115 p.Google Scholar
Campbell, K. E. Jr. 1982. Late Pleistocene events along the coastal plain of northern South America, p. 423440. In Prance, G. T. (ed.), Biological Diversification in the Tropics. Columbia University Press, New York.Google Scholar
Campbell, K. E. Jr., Frailey, C. A., and Romero Pittman, L. 2000. The late Miocene gomphothere Amahuacatherium peruvium (Proboscidea: Gomphotheriidae) from Amazonian Peru: Implications for the Great American Faunal Interchange. Boletín del Instituto Geológico Minero y Metalúrgico, Serie D: Estudios Regionales, 23:1152.Google Scholar
Carranza-Castañeda, O. and Miller, W. E. 2004. Late Tertiary terrestrial mammals from central Mexico and their relationship to South American immigrants. Revista Brasileira de Paleontología, 7(2):249261.Google Scholar
Cartelle, C. 1999. Pleistocene mammals of the Cerrado and Caatinga of Brazil, p. 2746. In Eisenberg, J. F. and Redford, R. K. (eds.), Mammals of the Neotropics. Volume 3. The Central Neotropics: Ecuador, Peru, Bolivia, Brazil. University of Chicago Press, Chicago.Google Scholar
Cartelle, C. and Langguth, A. 1999. Protocyon troglodytes (Lund): Um canídeo intertropical extinto. Anais da Academia Brasileira do Ciências, 71:371384.Google Scholar
Churcher, C. S. 1959. Fossil canids from tar pits of La Brea, Peru. Science, 130:364365.Google Scholar
Cione, A. L. and Tonni, E. P. 2001. Correlation of Pliocene to Holocene southern South America and European vertebrate-bearing units. Bolletino della Societá Paleontologica Italiana, 40:167173.Google Scholar
Clutton-Brock, J., Corbet, G. B., and Hills, M. 1976. A review of the family Canidae, with a classification by numerical methods. Bulletin of the British Museum (Natural History), Zoology, 29:119199.Google Scholar
Coates, A. G. 2003. La forja de Centroamérica, p. 140. In Coates, A. G. (ed.), Paseo Pantera. Una Historia de la Naturaleza y Cultura de Centroamérica. Smithsonian Istitution Press, Washington, DC.Google Scholar
Coates, A. G. and Obando, J. A. 1996. The geologic evidence of the Central American Isthmus, p. 2156. In Jackson, B. C., Budd, A. F., and Coates, A. G. (eds.), Evolution and Environment in Tropical America. The University of Chicago Press, Chicago.Google Scholar
Colinvaux, P. A. 1996. Quaternary environmental history and forest diversity in the Neotropics, p. 359406. In Jackson, B. C., Budd, A. F., and Coates, A. G. (eds.), Evolution and Environment in Tropical America. The University of Chicago Press, Chicago.Google Scholar
Colinvaux, P. A. 2003. La historia de los bosques del Istmo desde la era glacial hasta el presente, p. 137152. In Coates, A. G. (ed.), Paseo Pantera. Una Historia de la Naturaleza y Cultura de Centroamérica. Smithsonian Istitution Press, Washington, DC.Google Scholar
Czaplewski, N. J., Rincón, A. D., and Morgan, G. S. 2005. Fossil bats (Mammalia, Chiroptera) remains from Inciarte Tar Pit, Sierra de Perijá, Venezuela. Caribean Journal of Science, 41(4):768781.Google Scholar
Desmaret, A. G. 1822. Mammalogie ou Description des Espéces de Mammiféres. 2. Agasse, Paris, 555 p.Google Scholar
Dundas, R. G. 1999. Quaternary records of the dire wolf, Canis dirus, in North and South America. Boreas, 28:375385.Google Scholar
Eisenberg, J. F. 1989. Mammals of the Neotropics. Volume 1. The Northern Neotropics: Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana. University of Chicago Press, Chicago, 624 p.Google Scholar
Evans, H. E. 1993. Miller's Anatomy of the Dog, (third edition). W. B. Saunders Company, Philadelphia, 1113 p.Google Scholar
Fischer, G. 1814. Zoognosia Tabulis Synopticis Illustrate. Volumen tertium. Quadrupedum reliquorum, cetorum et montrymatum descriptionem continens. Nicolai Sergeidis Vsevolozsky, Mosquaea, 732 p.Google Scholar
Fischer, G. 1817. Adversaria Zoologica. Mémoire Société Impériale Naturelle, 5:368428.Google Scholar
Flynn, J. J., Kowallis, B. J., Nuñez, C., Carranza-Castañeda, O., Miller, W. E., Swisher, C. C. III, and Lindsay, E. 2005. Geochronology of Hemphillian-Blancan aged strata, and implications for the timing of the Great American Biotic Interchange. Journal of Geology, 113:287307.Google Scholar
Fritzell, E. K. and Haroldson, K. J. 1982. Urocyon cinereoargenteus . Mammalian Species, 189:18.Google Scholar
Fuller, T. K. and Cypher, B. L. 2004. Urocyon cinereoargenteus (Schreber, 1775), p. 9297. In Sillero Zubiri, C., Hoffman, M., and Macdonald, D. W. (eds.), Canids: Foxes, Wolves, Jackals and Dogs. Status Survey and Conservation Plan. IUCN, Gland.Google Scholar
Getz, L. L. 1960. Middle Pleistocene carnivores from southwestern Kansas. Journal of Mammalogy, 41:361365.Google Scholar
Giebel, C. G. 1855. Die Saugethiere in Zoologisher, Anatomischer, und Palantologoscher Beziehung Umfassend Dargetellt. Abel, Leipzig, 1108 p.Google Scholar
Graham, R. W. and Lundelius, E. L. 1994. FAUNMAP: A database documenting Late Quaternary distributions of mammal species in the United States. Illinois State Museum Scientific Paper, 25:1690.Google Scholar
Hall, E. R. and Kelson, K. R. 1959. The Mammals of North America. Ronald Press Company, New York, 536 p.Google Scholar
Hamilton Smith, C. 1839. Dogs. Canidae or genus Canis of authors. Volume 1. Jardine's Naturalist Library, 9:1267.Google Scholar
Hershkovitz, P. 1972. The recent mammals of the Neotropical Region: A zoogeographic and ecological review, p. 311431. In Keast, A., Erk, F. C., and Glass, B. (eds.), Evolution, Mammals, and Southern Continents. State University of New York Press, Albany.Google Scholar
Hodgson, B. H. 1833. Description of the wild dog of the Himalaya (Canis primaevus). Asiatick Researches, 18:221237.Google Scholar
Hoffstetter, R. 1952. Les mammiféres pléistocénes de la République de l'Equateur. Société Géologique de France Mémoirs (n.s.), 31(66):1391.Google Scholar
Hoffstetter, R. 1963. La faune Pleistocéne de Tarija (Bolivie), note preliminaire. Bulletin Muséum National d' Histoire Naturelle, 35:194203.Google Scholar
Hoffstetter, R. 1986. High Andean mammalian faunas during the Plio-Pleistocene, p. 218245. In Vuilleumier, F., and Monasterds, M. (eds.), High Altitude Sub Tropical Biogeography. Oxford University Press, Oxford.Google Scholar
Illiger, K. 1804–1811 [1815]. Überlick der Säugethiere nach ihrer Verteitung über die Welttheile. Akademie der Wissenshaften Berlin Abhandlungen, 1804–1811:39159.Google Scholar
Jull, A. T. L., Iturralde-Vinent, M., O'malley, J. M., Macphee, R. D. E., Mcdonald, H. G., Martin, P. S., Moody, J., and Rincón, A. 2004. Radiocarbon dating of extinct fauna in the Americas recovered from tar pits. Nuclear Instruments and Methods in Physics Research B, 223–224:668671.Google Scholar
Kirby, M. X. and Macfadden, B. J. 2005. Was southern Central America an archipelago or a peninsula in the middle Miocene? A test using land-mammal body size. Palaeogeography, Palaeoclimatology, Palaeoecology, 228(3–4):173202.Google Scholar
Kraglievich, J. L. 1952. El perfil geológico de Chapadmalal y Miramar. Provincia de Buenos Aires. Revista del Museo de Ciencias Naturales y Tradicional de Mar del Plata, 1:837.Google Scholar
Kraglievich, L. 1928. Contribución al conocimiento de los grandes cánidos extinguidos de Sud América. Anales de la Sociedad Científica Argentina, 106:2566.Google Scholar
Kraglievich, L. 1930. Craneometría y clasificación de los cánidos sudamericanos, especialmente los argentinos actuales y fósiles. Physis, 10:3573.Google Scholar
Kretzoi, M. 1943. Kochitis centenii n. g. n. sp., ein altertumlicher Creodonte aus dem Oberoligozan Siebenburgens. Foldtany Kozlony, 52:10195.Google Scholar
Kunth, C. S. 1822. Malvaceae, Büttneriaceae, Tiliaceae: Familiae denuo ad examen revocatae characteribusque magis exactis distinctae. Nova Genera et Species Plantarum, 5:149.Google Scholar
Kurtén, B. 1984. Geographic differentiation in the Rancholabrean Dire Wolf (Canis dirus Leidy) in North America. Special Publication of the Carnegie Museum of Natural History, 8:218227.Google Scholar
Kurtén, B. and Anderson, E. 1980. Pleistocene Mammals of North America. Columbia University Press, New York, 442 p.Google Scholar
Langguth, A. 1970. Una nueva clasificación de los cánidos sudamericanos. Abstracts 4th Congreso Latinoamericano de Zoología, Montevideo, 1:129143.Google Scholar
Leidy, J. 1856. Description of some remains of extinct mammalia. Journal of the Academy National of Science, Philadelphia, ser. 2, 3:166171.Google Scholar
Leidy, J. 1858. Notice of remains of extinct Vertebrata, from the Valley of the Niobrara River, collected during the exploring expedition of 1857, in Nebraska, under the command of Lieut. G. K. Warren, U. S. Top. Eng., by Dr. F. V. Hayden, geologist of the Expedition. Proceedings of the Academy of Natural Sciences, 1858:2029.Google Scholar
Leidy, J. 1869. The extinct mammalian fauna of Dakota and Nebraska. Journal of the Academy National of Science, Philadelphia, ser. 2, 7:1472.Google Scholar
Lemon, R. R. H. and Churcher, C. S. 1961. Pleistocene geology and paleontology of the Talara region, Northwest Peru. American Journal of Science, 259:410429.Google Scholar
Linares, O. J. 1998. Mamíferos de Venezuela. Sociedad Conservacionista Audubon de Venezuela, Caracas, 691 p.Google Scholar
Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, Secumdum Classes, Ordines, Genera, Species cum Characteribus, Differentiis, Synonymis, Locis. Volume 1, (tenth edition). L. Salvii, Uppsala, 824 p.Google Scholar
Lund, P. 1837. Om huler I lalksteen I det indre af Brasilien, der tildeel indeholde fossile knokler. Anden Afhandling. Lappa da Cerca Grande. Det kongelige Danske Vindenskabernes Selskabs naturvidenskabelige og mathematiske Afhandliger, 6:307332.Google Scholar
Lund, P. W. 1838. Blik paa Brasiliens Dyreverden för Sidste Jordomvaeltning. Förste Afhandling: Indledning. Det Kongelige Danske Videnskabernes Selskabs Naturnvidenskabelige og Mathematiske Afhandlinger, 8:2560.Google Scholar
Lund, P. W. 1843. Blik paa Brasiliens Dyreverden för Sidste Jordomvaeltning. 5. Afhandling: Fortsaettelse af Pattedyrene. On de nulevende og udöde arter af rovdyrenes familia. Det kongelige Danske Vindenskabernes Selskabs naturvidenskabelige og mathematiske Afhandliger, 11:182.Google Scholar
Lund, P. W. 1950. Memórias Sôbre a Paleontologia Brasileira. Revistas e Comentadas por Carlos de Paula Couto. Ministerio da Educaçâo e Salude, Instituto Nacional do Livro, Rio de Janeiro, 589 p.Google Scholar
Macfadden, B. J. 2000. Middle Pleistocene climate change recorded in fossil mammal teeth from Tarija, Bolivia, and upper limit of the Ensenadan land-mammal age. Quaternary Research, 54:121131.Google Scholar
Martin, R. A. 1974. Fossil mammals from the Coleman IIA Fauna, Sumter County, p. 3598. In Webb, S. D. (ed.), Pleistocene Mammals of Florida. University Press of Florida, Gainesville.Google Scholar
Mcdonald, G. H., Moody, J. M., and Rincón, A. 1999. Preliminary report on Pleistocene vertebrates from asphalt deposits in the Maracaibo basin, Venezuela. Abstracts of the Congreso International: Evolución Neotropical del Cenozoico, La Paz, 1:27.Google Scholar
Mercerat, A. 1891. Caracteres diagnósticos de algunas especies de Creodonta conservadas en el Museo de La Plata. Revista del Museo de La Plata, 2:5152.Google Scholar
Merriam, J. C. 1912. The fauna of Rancho La Brea, Pt. II, Canidae. Memoirs of the University of California, 1:217273.Google Scholar
Merriam, J. C. 1918. Note of the systematic position of wolves of the Canis dirus group. University of California Publications, Bulletin, Department of Geology, 10(27):531533.Google Scholar
Mivart, Saint G. 1890. Dogs, Jackals, Wolves and Foxes. A Monograph of the Canidae. R. H. Porter, London, 216 p.CrossRefGoogle Scholar
Mondolfi, E. and Bisbal, F. 1985. Notas sobre el ‘Zorro Gris’ venezolano (Urocyon cinereoargenteus venezuelae, Allen, 1911) Mammalia, Carnívora. Acta Biológica Venezuelica, 12:918.Google Scholar
Moore, C. M. and Collins, P. W. 1995. Urocyon littoralis . Mammalian Species, 489:17.Google Scholar
Morgan, G. S. 2002. Late Rancholabrean mammals from southernmost Florida and Neotropical influence in Florida Pleistocene Faunas. Smithsonian Contributions to Paleobiology, 93:1538.Google Scholar
Munthe, K. 1998. Canidae, p. 124143. In Janis, C., Scott, K. M., and Jacobs, L. L. (eds.), Evolution of Tertiary Mammals of North America. Volume 1. Cambridge University Press, Cambridge.Google Scholar
Nigra, J. O. and Lance, J. F. 1947. A statistical study of the metapodials of the dire wolf group from the Pleistocene of Rancho La Brea. Bulletin Southern California Academy of Sciences, 46:2634.Google Scholar
Nowak, R. M. 1979. North American Quaternary Canis . University of Kansas Museum of Natural History Monograph, 6, 154 p.Google Scholar
Nowak, R. M. 2002. The original status of wolves in eastern North America. Southern Naturalist, 1:95130.Google Scholar
Prevosti, F. J., Dondas, A., and Isla, F. I. 2004. Revisión del registro de Theriodictis Mercerat, 1891 (Carnivora, Canidae) y descripción de un nuevo ejemplar de Theriodictis platensis Mercerat, 1891 del Pleistoceno de la provincia de Buenos Aires (Argentina). Ameghiniana, 41:245250.Google Scholar
Prevosti, F. J., Zurita, A., and Carlini, A. 2005. Biostratigraphy, systematics and palaeoecology of the species of Protocyon Giebel, 1855 (Carnivora, Canidae) in South America. Journal of South American Earth Science, 20:512.Google Scholar
Rincón, R. A. D. 2005. Los roedores fósiles presentes en el Mene de Inciarte, Sierra de Perijá, estado Zulia, Venezuela. Bioestratigrafía e implicaciones paleoambientales. Unpublished doctoral dissertation, Instituto Venezolano de Investigaciones Científicas (IVIC), Caracas, 274 p.Google Scholar
Royo Y Gómez, J. 1960. El yacimiento de vertebrados pleistocenos de Muaco, Estado Falcón, con industria lítica humana. Abstracts of the Internacional Geological Congress, Copenhague, 21:154157.Google Scholar
Schreber, J. C. D. 1775. Die Säugethiere in Abbildungen Nach der Nature mit Beschreibungen. 3. W. Walther, Leipzig, 590 p.Google Scholar
Schubert, C. 1989. Paleoclima del Pleistoceno Tardío en el Caribe y regiones adyacentes: Un intento de compilación. Ciencias de la Tierra y el Espacio, 15–16:4058.Google Scholar
Sellards, E. H. 1916. Human remains and associated fossils from the Pleistocene of Florida. Annual Report of the Florida Geological Survey, 8:123160.Google Scholar
Simpson, G. G. 1929. Pleistocene mammalian fauna of the Seminole Field, Pinellas County, Florida. Bulletin of the American Museum of Natural History, 56:561599.Google Scholar
Smith, J. B. and Dodson, P. 2003. A proposal for a standard terminology of anatomical notation and orientation in fossil vertebrate dentitions. Journal of Vertebrate Paleontology, 23:112.Google Scholar
Steenkamp, G. and Borrel, C. G. 1999. Oral and dental conditions in adult African wild dog skull: A preliminary report. Journal of Veterinary Dentistry, 16:6568.Google Scholar
Stevens, M. S. 1965. A new species of Urocyon from the Upper Pliocene of Kansas. Journal of Mammalogy, 46:265269.Google Scholar
Stock, C. and Lance, J. F. 1948. The relative lengths of limb elements in Canis dirus . Bulletin Southern California Academy of Sciences, 47:7984.Google Scholar
Szuma, E. 2002. Dental polymorphism in a population of the red fox (Vulpes vulpes) from Poland. Journal of Zoology London, 256:243253.Google Scholar
Tedford, R. H., Taylor, B. E., and Wang, X. 1995. Phylogeny of the Caninae (Carnivore: Canidae): The living taxa. American Museum Novitates, 3146:137.Google Scholar
Urbani, F. and Galarraga, F. 1991. Inventario de Menes de la Sierra de Perijá. Universidad Central de Venezuela. Facultad de Ingeniería. Escuela de Geología, Minas y Geofísica. Informe interno, 80 p.Google Scholar
Wang, X. 1993. Transformation from the plantigrady to digitigrady: Functional morphology of locomotion in Hesperocyon (Canidae: Carnivora). American Museum Novitates, 3069:123.Google Scholar
Wang, X. 1994. Phylogenetics systematics of the Hesperocyoninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History, 221:1207.Google Scholar
Wang, X., Tedford, R. H., and Taylor, B. E. 1999. Phylogenetics systematics of the Borophaginae (Carnivora: Canidae). Bulletin of the American Museum of Natural History, 243:1391.Google Scholar
Wang, X., Tedford, R. H., Van Valkenburgh, B., and Wayne, R. K. 2004. Phylogeny, classification, and evolutionary ecology of the Canidae, p. 820. In Sillero Zubiri, C., Hoffman, M., and Macdonald, D. W. (eds.), Canids: Foxes, Wolves, Jackals and Dogs. Status Survey and Conservation Plan. IUCN, Gland.Google Scholar
Wayne, R. K., George, S. B., Gilbert, D., Collins, P. W., Kovach, S. D., Girman, D., and Lehman, N. 1991. A morphological and genetic study of the Island Fox, Urocyon littoralis . Evolution, 45:18491868.Google Scholar
Webb, S. D. 1991. Ecogeography and the Great American Interchange. Paleobiology, 17:266280.Google Scholar
Webb, S. D. 1999. Isolation and interchange. A deep history of South American mammals, p. 1326. In Eisemberg, J. F. and Redford, R. K. (eds.), Mammals of the Neotropics. Volume 3. The Central Neotropics: Ecuador, Peru, Bolivia, Brazil. University of Chicago Press, Chicago.Google Scholar
Webb, S. D. 2003. El Gran Intercambio Biotico Americano, p. 107136. In Coates, A. G. (ed.), Paseo Pantera. Una Historia de la Naturaleza y Cultura de Centroamérica. Smithsonian Istitution Press, Washington, DC.Google Scholar
Webb, S. D. and Rancy, A. 1996. Late Cenozoic evolution of the Neotropical mammals, p. 335358. In Jackson, B.C., Budd, A. F., and Coates, A. G. (eds.), Evolution and Environment in Tropical America. The University of Chicago Press, Chicago.Google Scholar
Winge, H. 1895. Jordfunde og nulevende Rovdyr (Carnivora) fra Lagoa Santa, Minas Geraes, Brasilien. E Museo Lundii, 2(4): 1103.Google Scholar
Wozencraft, W. C. 1993. Order Carnivora, p. 279348. In Wilson, D. E., and Reeder, D. A. M. (eds.), Mammals Species of the World. A taxonomic and Geographic Reference. Smithsonian Institution Press, Washington, DC.Google Scholar