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Strangers in a strange land: Ecological dissimilarity to metatherian carnivores may partly explain early colonization of South America by Cyonasua-group procyonids

  • Russell K. Engelman (a1) and Darin A. Croft (a2)


It was once thought that the endemic carnivorous mammals of South America, the metatherian sparassodonts, were driven extinct by North American carnivorans through competitive exclusion. However, sparassodonts went extinct before most groups of carnivorans entered South America; only the endemic Cyonasua-group procyonids (Cyonasua and Chapalmalania), which immigrated to South America nearly 4 million years earlier than other carnivorans, significantly overlapped with sparassodonts in time. In this study, we examine the functional morphology of the dentition of Cyonasua and Chapalmalania through quantitative analysis to determine the dietary habits of these taxa and the degree to which they may have ecologically overlapped sparassodonts and large predatory Neogene didelphimorphians. We find Cyonasua and Chapalmalania to be more carnivorous than extant procyonids, other than Bassariscus, in agreement with previous studies, but more omnivorous than most other carnivorans and all meat-eating South American metatherians, including sparassodonts. The extreme ecological dissimilarity between Cyonasua-group procyonids and members of the endemic South American predator guild may explain why procyonids were able to successfully establish themselves in South America several million years earlier than most other northern mammals (including all other carnivorans): they moved into a previously unoccupied ecological niche (large omnivore) and avoided direct competition with incumbent native species, a situation similar to that documented in historical cases of biological invasion. The omnivorous diets and climbing/swimming abilities of procyonids may have increased their chances for a successful over-water dispersal relative to other carnivorans, further favoring their successful establishment in South America.



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Alvarenga, H., Chiappe, L., and Bertelli, S.. 2011. Phorusrhacids: the terror birds. Pp. 187208 in Dyke, G. and Kaiser, G., eds. Living dinosaurs: the evolutionary history of modern birds. Wiley, Oxford, U.K.
Ameghino, C., and Kraglievich, J.. 1925. Un nuevo prociónido cercoleptoide en el Pampeano inferior de la Argentina, Brachynasua meranii, n. gen., n. sp. Comunicaciones Museo Nacional de Historia Natural, “Bernadino Rivadavia,” Buenos Aires 2:181–191.
Azzurro, E., Tuset, V. M., Lombarte, A., Maynou, F., Simberloff, D., Rodríguez-Pérez, A., and Solé, R. V.. 2014. External morphology explains the success of biological invasions. Ecology Letters 17:14551463.
Baskin, J. A. 1982. Tertiary Procyoninae (Mammalia: Carnivora) of North America. Journal of Vertebrate Paleontology 2:7193.
Baskin, J. A. 2003. New procyonines from the Hemingfordian and Barstovian of the Gulf Coast and Nevada, including the first fossil record of the Potosini. Bulletin of the American Museum of Natural History 279:125146.
Baskin, J. A. 2004. Bassariscus and Probassariscus (Mammalia, Carnivora, Procyonidae) from the early Barstovian (middle Miocene). Journal of Vertebrate Paleontology 24:709720.
Beck, R. M. D., and Taglioretti, M. L.. 2019. A nearly complete juvenile skull of the marsupial Sparassocynus derivatus from the Pliocene of Argentina, the affinities of “sparassocynids,” and the diversification of opossums (Marsupialia; Didelphimorphia; Didelphidae). Journal of Mammalian Evolution doi: 10.1007/s10914-019-09471-y.
Berman, W. D. 1994. Los carnívoros continentales (Mammalia, Carnivora) del Cenozoico en la provincia de Buenos Aires. Universidad Nacional de La Plata, La Plata, Argentina.
Bigler, W. J., Hoff, G. L., and Johnson, A. S.. 1981. Population characteristics of Procyon lotor marinus in estuarine mangrove swamps of southern Florida. Florida Scientist 44:151157.
Burgin, C. J., Colella, J. P., Kahn, P. L., and Upham, N. S.. 2018. How many species of mammals are there? Journal of Mammalogy 99:114.
Creel, S., and Creel, N. M.. 1996. Limitation of African wild dogs by competition with larger carnivores. Conservation Biology 10:526538.
Croft, D. A., Engelman, R. K., Dolgushina, T., and Wesley, G.. 2018. Diversity and disparity of sparassodonts (Metatheria) reveal non-analogue nature of ancient South American mammalian carnivore guilds. Proceedings of the Royal Society of London B 285:20172012.
Cuaron, A. D., Valenzuela-Galvan, D., Garcia-Vasco, D., Copa, M. E., Bautista, S., Mena, H., Martinez-Godinez, D., Gonzalez-Baca, C., Bojorquez-Tapia, L. A., Barraza, L., Grammont, de, P. C., Galindo-Maldonado, F., Martinez-Morales, M. A., Vazquez-Dominguez, E., Andresen, E., Benitez-Malvido, J., Perez-Salicrup, D., McFadden, K. W., and Gompper, M. E.. 2009. Conservation of the endemic dwarf carnivores of Cozumel Island, Mexico. Small Carnivore Conservation 41:1521.
Czaplewski, N. J. 1996. Opossums (Didelphidae) and bats (Noctilionidae and Molossidae) from the late Miocene of the Amazon Basin. Journal of Mammalogy 77:8494.
Degrange, F. J., Noriega, J. I., and Areta, J. I.. 2012. Diversity and paleobiology of Santacrucian birds. Pp. 138155 in Vizcaíno, S. F., Kay, R. F., and Bargo, M. S., eds. Early Miocene paleobiology in Patagonia: high-latitude paleocommunities of the Santa Cruz Formation. Cambridge University Press, Cambridge.
Eizirik, E. 2012. A molecular view on the evolutionary history and biogeography of Neotropical carnivores (Mammalia, Carnivora). Pp. 123142 in Patterson, B. D. and Costa, L. P., eds. Bones, clones, and biomes. the history and geography of recent Neotropical mammals. University of Chicago Press, Chicago.
Engelman, R. K., and Croft, D. A.. 2014. A new species of small-bodied sparassodont (Mammalia, Metatheria) from the middle Miocene locality of Quebrada Honda, Bolivia. Journal of Vertebrate Paleontology 34:672688.
Flannery Sutherland, J. T., Moon, B. C., Stubbs, T. L., and Benton, M. J.. 2019. Does exceptional preservation distort our view of disparity in the fossil record? Proceedings of the Royal Society of London B 286:20190091.
Fleagle, J. G., Kay, R. F., and Anthony, M. R. L.. 1997. Fossil New World monkeys. Pp. 473495 in Kay, R. F., Madden, R. H., Cifelli, R. L., and Flynn, J. J., eds. Vertebrate paleontology in the Neotropics: the Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington, D.C.
Flynn, J. J., Guerrero, J., and Swisher, C. C. III. 1997. Geochronology of the Honda Group. Pp. 4459 in Kay, R. F., Madden, R. H., Cifelli, R. L., and Flynn, J. J., eds. Vertebrate paleontology in the Neotropics: the Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington, DC.
Forasiepi, A. M., Martinelli, A. G., and Goin, F. J.. 2007. Revisión taxonómica de Parahyaenodon argentinus Ameghino y sus implicancias en el conocimiento de los grandes mamíferos carnívoros del Mio-Plioceno de América de Sur. Ameghiniana 44:143159.
Forasiepi, A. M., Soibelzon, L. H., Gomez, C. S., Sánchez, R., Quiroz, L. I., Jaramillo, C., and Sánchez-Villagra, M. R.. 2014. Carnivorans at the Great American Biotic Interchange: new discoveries from the northern Neotropics. Naturwissenschaften 101:965974.
Gingerich, P. D., and Smith, B. H.. 1985. Allometric scaling in the dentition of primates and insectivores. Pp. 257272 in Jungers, W. L., ed. Size and scaling in primate biology. Springer US, Boston, Mass.
Goin, F. J. 1997. New clues for understanding Neogene marsupial radiations. Pp. 187206 in Kay, R. F., Madden, R. H., Cifelli, R. L., and Flynn, J. J., eds. Vertebrate paleontology in the Neotropics: the Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington, D.C.
Gompper, M. E. 1997. Population ecology of the white-nosed coati (Nasua narica) on Barro Colorado Island, Panama. Journal of Zoology 241:441455.
Hammer, Ø., Harper, D. A. T., and Ryan, P. D.. 2001. PAST: paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4:19.
Helgen, K. M., and Wilson, D. E.. 2005. A systematic and zoogeographic overview of the raccoons of Mexico and Central America. Pp. 221236 in Sánchez-Cordero, V. and Medellín, R. A., eds. Contribuciones Mastozoologicas en Homenaje a Bernardo Villa. Instituto de Biología, Universidad Nacional Autónoma de México, Mexico City.
Hutchinson, G. E. 1959. Homage to Santa Rosalia or why are there so many kinds of animals? American Naturalist 93:145159.
Kay, R. F. 1975. The functional adaptations of primate molar teeth. American Journal of Physical Anthropology 43:195215.
Kay, R. F., Madden, R. H., Cifelli, R. L., and Flynn, J. J., eds. 1997. Vertebrate paleontology in the Neotropics: the Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington, D.C.
Koepfli, K.-P., Gompper, M. E., Eizirik, E., Ho, C.-C., Linden, L., Maldonado, J. E., and Wayne, R. K.. 2007. Phylogeny of the Procyonidae (Mammalia: Carnivora): molecules, morphology and the Great American Interchange. Molecular Phylogenetics and Evolution 43:10761095.
Kraglievich, J. L., and de Olazabal, A. G.. 1959. Los prociónidos extinguidos del género Chapalmalania Amegh. Revista del Museo Argentino de Ciencias Naturales, Ciencias Zoologícas 6:159.
Lindqvist, C., Schuster, S. C., Sun, Y., Talbot, S. L., Qi, J., Ratan, A., Tomsho, L. P., Kasson, L., Zeyl, E., Aars, J., Miller, W., Ingólfsson, Ó., Bachmann, L., and Wiig, Ø.. 2010. Complete mitochondrial genome of a Pleistocene jawbone unveils the origin of polar bear. Proceedings of the National Academy of Sciences USA 107:50535057
Linnell, J. D. C., and Strand, O.. 2000. Interference interactions, co-existence and conservation of mammalian carnivores. Diversity and Distributions 6:169176.
López-Aguirre, C., Archer, M., Hand, S. J., and Laffan, S. W.. 2017. Extinction of South American sparassodontans (Metatheria): environmental fluctuations or complex ecological processes? Palaeontology 60:91115.
Lyras, G. A., van der Geer, A. A. E., and Rook, L.. 2010. Body size of insular carnivores: evidence from the fossil record. Journal of Biogeography 37:10071021.
Marshall, L. G. 1976. Evolution of the Thylacosmilidae, extinct saber-tooth marsupials of South America. Paleobios 23:130.
Marshall, L. G. 1977. Evolution of the carnivorous adaptive zone in South America. Pp. 709721 in Hecht, M. K., Goody, P. C., and Hecht, B. M., eds. Major patterns in vertebrate evolution. Plenum, New York.
Marshall, L. G. 1978. Evolution of the Borhyaenidae, extinct South American predaceous marsupials. University of California Publications in Geological Sciences 117:189.
Marshall, L. G. 1979. Review of the Prothylacyninae, an extinct subfamily of South American “dog-like” marsupials. Fieldiana (Geology), new series 3:149.
Marshall, L. G. 1981. The Great American Interchange: an invasion induced crisis for South American mammals. Pp. 133229 in Nitecki, M. H., ed. Biotic crises in ecological and evolutionary time. Academic Press, New York.
Marshall, L. G., Butler, R. F., Drake, R. E., Curtis, G. H., and Tedford, R. H.. 1979. Calibration of the Great American Interchange. Science 204:272279.
McFadden, K. W., Gompper, M. E., Valenzuela, D. G., and Morales, J. C.. 2008. Evolutionary history of the critically endangered Cozumel dwarf carnivores inferred from mitochondrial DNA analyses. Journal of Zoology 276:176186.
Meiri, S., Dayan, T., and Simberloff, D.. 2004. Body size of insular carnivores: little support for the island rule. American Naturalist 163:469479.
Mitchell, K. J., Bray, S. C., Bover, P., Soibelzon, L., Schubert, B. W., Prevosti, F., Prieto, A., Martin, F., Austin, J. J., and Cooper, A.. 2016. Ancient mitochondrial DNA reveals convergent evolution of giant short-faced bears (Tremarctinae) in North and South America. Biology Letters 12:20160062.
Montes, C., Cardona, A., Jaramillo, C., Pardo, A., Silva, J. C., Valencia, V., Ayala, C., Pérez-Angel, L. C., Rodriguez-Parra, L. A., Ramirez, V., and Niño, H.. 2015. Middle Miocene closure of the Central American Seaway. Science 348:226229.
O'Dea, A., Lessios, H. A., Coates, A. G., Eytan, R. I., Restrepo-Moreno, S. A., Cione, A. L., Collins, L. S., de Queiroz, A., Farris, D. W., Norris, R. D., Stallard, R. F., Woodburne, M. O., Aguilera, O., Aubry, M.-P., Berggren, W. A., Budd, A. F., Cozzuol, M. A., Coppard, S. E., Duque-Caro, H., Finnegan, S., Gasparini, G. M., Grossman, E. L., Johnson, K. G., Keigwin, L. D., Knowlton, N., Leigh, E. G., Leonard-Pingel, J. S., Marko, P. B., Pyenson, N. D., Rachello-Dolmen, P. G., Soibelzon, E., Soibelzon, L., Todd, J. A., Vermeij, G. J., and Jackson, J. B. C.. 2016. Formation of the Isthmus of Panama. Science Advances 2:e1600883.
Oldfield, C. C., McHenry, C. R., Clausen, P. D., Chamoli, U., Parr, W. C. H., Stynder, D. D., and Wroe, S.. 2011. Finite element analysis of ursid cranial mechanics and the prediction of feeding behaviour in the extinct giant Agriotherium africanum. Journal of Zoology 286:163170.
Orcutt, J. D. 2015. Ecomorphology of Australian carnivore guilds. Journal of Vertebrate Paleontology, Program and Abstracts, p. 190.
Ortiz-Ramírez, M. F., Sánchez-González, L. A., Castellanos-Morales, G., Ornelas, J. F., and Navarro-Sigüenza, A. G.. 2018. Concerted Pleistocene dispersal and genetic differentiation in passerine birds from the Tres Marías Archipelago, Mexico. The Auk 135:716732.
Palomares, F., and Caro, T. M.. 1999. Interspecific killing among mammalian carnivores. American Naturalist 153:492508.
Patterson, B., and Pascual, R.. 1968. The fossil mammal fauna of South America. Quarterly Review of Biology 43:409451.
Pearson, D. E., Ortega, Y. K., and Sears, S. J.. 2012. Darwin's naturalization hypothesis up-close: intermountain grassland invaders differ morphologically and phenologically from native community dominants. Biological Invasions 14:901913.
Pol, D., Leardi, J. M., Lecuona, A., and Krause, M.. 2012. Postcranial anatomy of Sebecus icaeorhinus (Crocodyliformes, Sebecidae) from the Eocene of Patagonia. Journal of Vertebrate Paleontology 32:328354.
Prevosti, F. J., Forasiepi, A., and Zimicz, N.. 2013. The evolution of the Cenozoic terrestrial mammal guild in South America: competition or replacement? Journal of Mammalian Evolution 20:321.
Prevosti, F. J., and Forasiepi, A. M.. 2018. Evolution of South American mammalian predators during the Cenozoic: palaeobiogeographic and palaeoenvironmental contengencies. Springer, Cham, Switzerland.
Prothero, D. R., Campbell, K. E., Beatty, B. L., and Frailey, C. D.. 2014. New late Miocene dromomerycine artiodactyl from the Amazon Basin: implications for interchange dynamics. Journal of Paleontology 88:434443.
Reguero, M. A., and Candela, A. M.. 2011. Late Cenozoic mammals from the northwest of Argentina. Pp. 411426 in Salfity, J. A. and Marquillas, R. A., eds. Cenozoic geology of the Central Andes of Argentina. SCS Publisher, Salta, Argentina.
Reyes, M. d. l., Poiré, D., Soibelzon, L., Zurita, A. E., and Arrouy, M. J.. 2013. First evidence of scavenging of a glyptodont (Mammalia, Glyptodontidae) from the Pliocene of the Pampean region (Argentina): taphonomic and paleoecological remarks. Palaeontologia Electronica 16:15A: 113.
Rick, T. C., Erlandson, J. M., Vellanoweth, R. L., Braje, T. J., Collins, P. W., Guthrie, D. A., and Stafford, T. W.. 2009. Origins and antiquity of the island fox (Urocyon littoralis) on California's Channel Islands. Quaternary Research 71:9398.
Riggs, E. S. 1934. A new marsupial saber-tooth from the Pliocene of Argentina and its relationships to other South American predacious marsupials. Transactions of the American Philosophical Society 24:132.
Robinson, J. G., and Redford, K. H.. 1986. Body size, diet, and population density of Neotropical forest mammals. American Naturalist 128:665680.
Rodriguez, S. G., Soibelzon, L. H., Rodrigues, S., Morgan, C. C., Bernardes, C., Avilla, L., and Lynch, E.. 2013. First record of Procyon cancrivorus (G. Cuvier, 1798) (Carnivora, Procyonidae) in stratigraphic context in the Late Pleistocene of Brazil. Journal of South American Earth Sciences 45:15.
Ruiz-Ramoni, D., Rincón, A., and Montellano-Ballesteros, M.. 2019. Evidencias del origen de Nasua y Procyon (Procyonidae: Carnivora) en América del Sur. Revista Brasileira de Paleontologia 21:8794.
Sacco, T., and Van Valkenburgh, B. 2004. Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae). Journal of Zoology 263:4154.
Scapino, R. 1981. Morphological investigation into functions of the jaw symphysis in carnivorans. Journal of Morphology 167:339–75.
Scott, J. E., Hogue, A. S., and Ravosa, M. J.. 2012. The adaptive significance of mandibular symphyseal fusion in mammals. Journal of Evolutionary Biology 25:661673.
Sih, A., Bolnick, D. I., Luttbeg, B., Orrock, J. L., Peacor, S. D., Pintor, L. M., Preisser, E., Rehage, J. S., and Vonesh, J. R.. 2010. Predator–prey naïveté, antipredator behavior, and the ecology of predator invasions. Oikos 119:610621.
Simpson, G. G. 1950. History of the fauna of Latin America. American Scientist 38:361389.
Simpson, G. G. 1980. Splendid isolation: the curious history of South American mammals. Yale University Press, New Haven, Conn.
Skóra, F., Abilhoa, V., Padial, A. A., and Vitule, J. R. S.. 2015. Darwin's hypotheses to explain colonization trends: evidence from a quasi-natural experiment and a new conceptual model. Diversity and Distributions 21:583594.
Soibelzon, E., Gasparini, G. M., Zurita, A. E., and Soibelzon, L. H.. 2008. Análisis faunístico de vertebrados de las toscas del Río de La Plata (Buenos Aires, Argentina): un yacimiento paleontológico en desaparición. Revista del Museo Argentino de Ciencias Naturales 10:291308.
Soibelzon, L., and Prevosti, F. J.. 2007. Los carnívoros (Carnivora, Mammalia) terrestres del Cuaternario de América del Sur. Pp. 4968 in Pons, G. X. and Vicens, D., eds. Geomorfología Litoral i Quaternari. Homenatge a Joan Cuerda Barceló. Monografia de la Societat d'Història Natural, Palma de Mallorca.
Soibelzon, L. H. 2011. First description of milk teeth of fossil South American procyonid from the lower Chapadmalalan (Late Miocene–Early Pliocene) of “Farola Monte Hermoso,” Argentina: paleoecological considerations. Paläontologische Zeitschrift 85:8389.
Soibelzon, L. H., Rinderknecht, A., Tarquini, J., and Ugalde, R.. 2019. First record of fossil procyonid (Mammalia, Carnivora) from Uruguay. Journal of South American Earth Sciences 92:368373.
Strait, S. G. 1993. Differences in occlusal morphology and molar size in frugivores and faunivores. Journal of Human Evolution 25:471484.
Tambussi, C. P., and Degrange, F. J.. 2013. The dominance of zoophagous birds: just a cliché? Pp. 87102 in Tambussi, C. P. and Degrange, F., eds. South American and Antarctic continental Cenozoic birds: paleobiogeographic affinities and disparities. Springer, Dordrecht, Netherlands.
Tarquini, J., Toledo, N., Morgan, C. C., and Soibelzon, L. H.. 2017. The forelimb of †Cyonasua sp. (Procyonidae, Carnivora): ecomorphological interpretation in the context of carnivorans. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 106:325335.
Tarquini, J., Toledo, N., Soibelzon, L. H., and Morgan, C. C.. 2018a. Body mass estimation for †Cyonasua (Procyonidae, Carnivora) and related taxa based on postcranial skeleton. Historical Biology 30:496506.
Tarquini, J., Vilchez Barral, M. G., and Soibelzon, L. H.. 2016. Los prociónidos fósiles de América del Sur. Pp. 359–365 in Agnolin, F. L., Lio, G. L., Brissón Egli, F., Chimento, N. R., and Novas, F. E., eds. Historia Evolutiva y Paleobiogeográfica de los Vertebrados de América del Sur. Museo Argentino de Ciencias Naturalis “Bernardino Rivadavia,” Buenos Aires.
Tarquini, S. D., Chemisquy, M. A., and Prevosti, F. J.. 2018b. Evolution of the carnassial in living mammalian carnivores (Carnivora, Didelphimorphia, Dasyuromorphia): Diet, phylogeny, and allometry. Journal of Mammalian Evolution. doi: 10.1007/s10914-018-9448-7.
Van der Geer, A. G., Vos de, Lyras, J., and Dermitzakis, M.. 2011. Evolution of island mammals: adaptation and extinction of placental mammals on islands. Wiley-Blackwell, Chichester, U.K.
Van Valkenburgh, B. 1990. Skeletal and dental predictors of body mass in carnivores. Pp. 181205 in Damuth, J. and MacFadden, B. J., eds. Body size in mammalian paleobiology: estimation and biological implications. Cambridge University Press, Cambridge.
Van Valkenburgh, B. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): evolutionary interactions among sympatric predators. Paleobiology 17:340362.
Vidal-García, M., and Keogh, J. S.. 2017. Invasive cane toads are unique in shape but overlap in ecological niche compared to Australian native frogs. Ecology and Evolution 7:76097619.
Vizcaíno, S. F., and De Iuliis, G.. 2003. Evidence for advanced carnivory in fossil armadillos (Mammalia: Xenarthra: Dasypodidae). Paleobiology 29:123138.
Vizcaíno, S. F., Fariña, R. A., Zárate, M. A., Bargo, M. S., and Schultz, P.. 2004. Palaeoecological implications of the mid-Pliocene faunal turnover in the Pampean Region (Argentina). Palaeogeography, Palaeoclimatology, Palaeoecology 213:101113.
Webb, S. D. 1985. Late Cenozoic mammal dispersals between the Americas. Pp. 357386 in Stehli, F. G. and Webb, S. D., eds. The Great American Biotic Interchange. Springer US, Boston, Mass.
Werdelin, L. 1987. Jaw geometry and molar morphology in marsupial carnivores; analysis of a constraint and its macroevolutionary consequences. Paleobiology 13:342350.
Werdelin, L., and Lewis, M. E.. 2013. Temporal change in functional richness and evenness in the eastern African Plio-Pleistocene carnivoran guild. PLoS ONE 8:e57944.
Werdelin, L., and Wesley-Hunt, G. D.. 2010. The biogeography of carnivore evolution. Pp. 225245 in Goswami, A. and Friscia, A., eds. Carnivoran evolution: new views on phylogeny, form, and function. Cambridge University Press, Cambridge.
Wesley-Hunt, G. D. 2005. The morphological diversification of carnivores in North America. Paleobiology 31:3555.
Woodburne, M. O. 2010. The Great American Biotic Interchange: dispersals, tectonics, climate, sea level and holding pens. Journal of Mammalian Evolution 17:245264.
Wroe, S., Argot, C., and Dickman, C.. 2004. On the rarity of big fierce carnivores and primacy of isolation and area: tracking large mammalian carnivore diversity on two isolated continents. Proceedings of the Royal Society of London B 271:12031211.
Zeveloff, S. I. 2002. Raccoons: a natural history. Smithsonian Institution Press, Washington, D.C.
Zimicz, N. 2014. Avoiding competition: the ecological history of late Cenozoic metatherian carnivores in South America. Journal of Mammalian Evolution 21:383393.


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