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Reassessment of the hairy long-nosed armadillo “Dasypus” pilosus (Xenarthra, Dasypodidae) and revalidation of the genus Cryptophractus Fitzinger, 1856



The hairy long-nosed armadillo, currently referred as Dasypus (Cryptophractus) pilosus, is an enigmatic species endemic to montane cloud forests and subparamo of Peruvian Andes. Its strikingly different external features, which include the carapace concealed by abundant hair, the presence of more movable bands, and a slender skull, have raised questions regarding its taxonomic status as subgenus or as genus. This paper assesses this issue based on a cladistic study and provides a detailed comparative description of the species, including the first account on the distinctive ornamentation of its osteoderms. Based on several unique characters in the carapace, skull, mandible, and teeth, as well as on the external phylogenetic position relative to other Dasypus, we favor the assignment of the hairy long-nosed armadillo to other genus. As result, we revalidate the original generic epithet, so that the valid name of the species is Cryptophractus pilosus Fitzinger, 1856.



Peru, Cingulata, Dasypodinae, woolly armadillo, Peruvian Andes

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Aguiar, J.M. & da Fonseca, G.A.B. (2008) Conservation status of the Xenarthra. In: Vizcaíno, S.F. & Loughry, W.J. (Eds.), The biology of the Xenarthra. University Press of Florida, Gainesville, pp. 215–231.

Benton, M.J. (2000) Stems, nodes, crown clades, and rank-free lists: is Linnaeus dead? Biological Reviews, 75 (4), 633–648.

Billet, G., Hautier, L., de Muizon, C. & Valentin, X. (2011) Oldest cingulate skulls provide congruence between morphological and molecular scenarios of armadillo evolution. Proceedings of the Royal Society B, 278, 2791–2797.

Burmeister, H. (1862) Beschreibung eines behaarten Gürtelthieres Praopus hirsutus, aus dem National-Museum zu Lima. Abhandlungen der Naturforschenden Gesellschaft zu Halle, 6, 145–148.

Cabrera, A. (1958) Catálogo de los mamíferos de América del Sur. Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Zoología, 4, 1–307.

Carlini, A.A., Vizcaíno, S.F. & Scillato-Yané, G.J. (1997) Armored xenarthrans: a unique taxonomic and ecologic assemblage. In: Kay, R.F., Madden, R.H., Cifelli, R.L. & Flynn, J.J (Eds.), Vertebrate paleontology in the Neotropics. The Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press, Washington/London, pp. 213–226.

Carlini, A.A., Castro, M.C., Madden, R.H. & Scillato-Yané, G.J. (2014) A new species of Dasypodidae (Xenarthra: Cingulata) from the late Miocene of northwestern South America: implications in the Dasypodini phylogeny and diversity. Historical Biology, 26 (6), 728–736.

Castro, M.C., Ribeiro, A.M., Ferigolo, J. & Langer, M.C. (2013a) Redescription of Dasypus punctatus Lund, 1840 and considerations on the genus Propraopus Ameghino, 1881 (Xenarthra, Cingulata). Journal of Vertebrate Paleontology, 33, 434–444.

Castro, M.C., Avilla, L.S., Freitas, M.L. & Carlini, A.A. (2013b) The armadillo Propraopus sulcatus (Mammalia: Xenarthra) from the late Quaternary of northern Brazil and a revised synonymy with Propraopus grandis. Quaternary International, 317, 80–87.

Castro, M.C., Carlini, A.A., Sánchez, R. & Sánchez-Villagra, M.R. (2014) A new Dasypodini armadillo (Xenarthra: Cingulata) from San Gregorio Formation, Pliocene of Venezuela: affinities and biogeographic interpretations. Naturwissenschaften, 101, 77–86.

Charles, C., Solé, F., Rodrigues, H.G. & Viriot, L. (2013) Under pressure? Dental adaptations to termitophagy and vermivory among mammals. Evolution, 67, 1792–1804.

Ciancio, M.R. (2010) Los Dasypodoidea (Mammalia, Xenarthra) del Deseadense (Oligoceno) de América del Sur. Su importancia filogenética y bioestratigráfica. Unpublished Doctoral Thesis. Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, 290 pp.

Ciancio, M.R., Vieytes, E.C., Castro, M.C. & Carlini, A.A. (2010) Estructura del esmalte en Dasypus (Xenarthra, Dasypodidae), consideraciones filogenéticas preliminares. XXIII Jornadas Argentinas de Mastozoología, Resúmenes, nº 26, 1–89.

Ciancio, M.R., Castro, M.C., Galliari, F.C., Carlini, A.A. & Asher, R.J. (2012) Evolutionary implications of dental eruption in Dasypus (Xenarthra). Journal of Mammalian Evolution, 19, 1–8.

Cope, E.D. (1889) The Edentata of North America. American Naturalist, 23, 657–664.

Costa, F.R. & Vizcaíno, S.F. (2010) A diagnostic character revisited: is there a fifth toe in the forefoot of Dasypus novemcinctus (Xenarthra: Dasypodidae)? Zootaxa, 2671, 61–64.

Delsuc, F., Superina, M., Tilak, M.K., Douzery, E.J.P. & Hassanin, A. (2012) Molecular phylogenetics unveils the ancient evolutionary origins of the enigmatic fairy armadillos. Molecular Phylogenetics and Evolution, 62: 673–680.

de Queiroz, K. & Donoghue, M.J. (2013) Phylogenetic nomenclature, hierarchical information, and testability. Systematic Biology, 62 (1), 167–174.

Engelmann, G.F. (1985) The phylogeny of the Xenarthra. In: Montgomery, G.G. (Ed.), The evolution and ecology of armadillos, sloths, and vermilinguas. Smithsonian Institution Press, Washington, pp. 51–63.

Feijó, A. & Cordeiro-Estrela, P. (2014) The correct name of the endemic Dasypus (Cingulata: Dasypodidae) from northwestern Argentina. Zootaxa, 3887 (1), 88–94.

Fitzinger, L.J. (1856) Versammlung Deutscher Naturforscher und Ärzte in Wien, Tageblatt, #32, 123.

Fitzinger, L.J. (1871) Die natürliche Familie der Gürtelthiere (Dasypodes). II. Abtheilung. Sitzungsberichte Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, 64, 329–390.

Flower, W.H. (1886) Exhibition of and remarks upon a rare species of armadillo belonging to the Museum of the Scarborough and Philosophical Society. Proceedings of the Zoological Society of London, 419–420.

Flower, W.H. (1894) Exhibition of and remarks upon a specimen of a Hairy Armadillo (Tatusia pilosa). Proceedings of the Zoological Society of London, 655.

Frechkop, S. & Yepes, J. (1949) Étude systématique et zoogéographique des dasypodidés conservés a l’Institut. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, 25 (5), 1–56.

Gaudin, T.J. (1999) The morphology of xenarthrous vertebrae (Mammalia, Xenarthra). Fieldiana, Geology, 41, 1–38.

Gaudin, T.J. & Wible, J.R. (2006) The phylogeny of living armadillos (Mammalia, Xenarthra, Cingulata): a craniodental analysis. In: Carrano, M.T., Gaudin, T.J., Blob, R.W. & Wible, J.R. (Eds.), Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds and Reptiles. The University of Chicago Press, Chicago, pp. 153–198.

Goloboff, P., Farris, J. & Nixon, K. (2008) TNT: a free program for phylogenetic analysis. Cladistics, 24, 774–786.

González-Ruiz, L.R., Ciancio, M.R. & Góis, F. (2013) El espécimen más completo de Peltephilidae Ameghino (Mammalia, Xenarthra, Cingulata): aportes sistemáticos. Ameghiniana, Abstract Supplement 50 (6), R41–R42.

Gray, J.E. (1821) On the natural arrangement of vertebrose animals. London Medical Repository, 15, 296–310.

Gray, J.E. (1865) Revision on the genera and species of Entomophagous, Edentata, founded on examination of the specimens in British Museum. Proceedings of the Zoological Society of London, 33 (1), 359–386.

Grimwood, I.R. (1969) Notes on the distribution and status of some Peruvian mammals 1968. American Commitee International Wild Life Protection and New York Zoological Society, Special Publication, 21, 1–86.

Hamlett, G.W.D. (1939) Identity of Dasypus septemcinctus Linnaeus with notes on some related species. Journal of Mammalogy, 20, 328–336.

Illiger, C. (1811) Prodromus systematis mammalium et avium additis terminis zoographicis utriudque classis. C. Salfeld, Berlin, 301 pp.

Jasinski, S.E. & Wallace, C. (2014) Investigation into the paleobiology of Dasypus bellus using geometric morphometrics and variation of the calcaneus. Journal of Mammalian Evolution, 21 (3), 285–298.

Kraft, R. (1995) Xenarthra. In: Niethammer, J., Schlieman, H. & Starck, D. (Eds.), Handbuch der Zoologie. Vol. 8. Mammalia. Verlag de Gruyter & Co., Berlin, pp. 1–79.

Kuntner, M. & Agnarsson, I. (2006) Are the Linnean and Phylogenetic Nomenclatural Systems Combinable? Recommendations for Biological Nomenclature. Systematic Biology, 55 (5), 774–784.

Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. 10a Edition. Laurentii Salvii, Stockholm, 824 pp.

Martin, B.E. (1916) Tooth development in Dasypus novemcinctus. Journal of Morphology, 27, 647–691.

McBee, K. & Baker, R.J. (1982) Dasypus novemcinctus. Mammalian Species, 162, 1–9.

Moeller, W. (1968) Allometrische Analyse der Gürteltierschädel. Ein Beitrag zur Phylogenie der Dasypodidae Bonaparte, 1838. Zoologische Jahrbücher Abteilung für Anatomie und Ontogenie der Tiere, 85, 411–528.

Mondolfi, E. (1967) Descripción de un nuevo armadillo del género Dasypus de Venezuela (Mammalia-Edentata). Memorias de la Sociedad de Ciencias Naturales La Salle, 78, 149–167.

NatureServe (2014) Predicted distribution of Dasypus pilosus. Available from: (Accessed 15 July 2014)

Pacheco, V. (2002) Mamíferos del Perú. In: Ceballos, G. & Simonetti, J. (Eds.), Diversidad y Conservación de los Mamíferos Neotropicales. CONABIO - UNAM. México, D.F., pp. 503–550.

Patterson, B., Segall, W. & Turnbull, W.D. (1989) The ear region in Xenarthrans (=Edentata: Mammalia). Part I, Cingulates. Fieldiana (Geology), 18, 1–46.

Smith, K.K. & Redford, K.H. (1990) The anatomy and function of the feeding apparatus in two armadillos (Dasypoda): anatomy is not destiny. Journal of Zoology, 222, 27–47.

Superina, M. & Abba, A.M. (2010) Dasypus pilosus. In: IUCN 2013. IUCN red list of threatened species. Version 2013.2. Available from: (22 January 2014)

Talmage, R.V. & Buchanan, G.D. (1954) The armadillo (Dasypus novemcinctus). A review of its natural history, ecology, anatomy and reproductive physiology. Rice Institute Pamphlet, Monograph in Biology, 41 (2), 1–135.

Thomas, O. (1927) The Godman-Thomas Expedition to Peru. V. Mammals collected by Mr. R. W. Hendee in the province of San Martin, N. Peru, mostly at Yurac Yacu. Annals and Magazine Natural History, Series 9, 19, 361–375.

Trouessart, E.-L. (1898) Catalogus mammalium tam viventium quam fossilium. Fasciculus V. Sirenia, Cetacea, Edentata, Marsupialia, Allotheria, Monotremata. Vol. 2. R. Friedländer & Sohn, Berolini, 266 pp. [pp. 999–1264]

Trouessart, E.-L. (1905) Catalogus mammalium tam viventium quam fossilium. Quinquennale supplementium (1899–1904). Cetacea, Edentata, Marsupialia, Allotheria, Monotremata.—Index alphabeticus. Vol. 4. R. Friedländer & Sohn, Berolini, 177 pp. [pp. 753–929]

Vizcaíno, S.F. (1994) Mecánica masticatoria de Stegotherium tesselatum Ameghino (Mammalia, Xenarthra) del Mioceno de Santa Cruz (Argentina). Algunos aspectos paleoecológicos relacionados. Ameghiniana, 31 (3), 283–290.

Vizcaíno, S.F. (1995) Identificación específica de las “mulitas”, género Dasypus L. (Mammalia, Dasypodidae), del noroeste argentino. Descripción de una nueva especie. Mastozoología Neotropical, 2 (1), 5–13.

Wetzel, R.M. (1982) Systematics, distribution, ecology, and conservation of South American edentates. In: Mares, M.A. & Genoways, H.H. (Eds.), Mammalian biology in South America. Special Publications Pymatuning Laboratory of Ecology, University of Pittsburgh, 6, pp. 345–375. [total page number 539 pp.]

Wetzel, R.M. (1985) Taxonomy and distribution of armadillos. In: Montgomery, G.G. (Ed.), The Evolution and Ecology of Armadillos, Sloths, and Vermilinguas. Smithsonian Institution, Washington D.C., pp. 23–46.

Wetzel, R.M. & Mondolfi, E. (1979) The subgenera and species of long-nosed armadillos, genus Dasypus. In: Eisenberg, J.F. (Ed.), Vertebrate Ecology in the Northern Neotropic. Smithsonian Institution Press, Washington D.C., pp. 43–63.

Wetzel, R.M., Gardner, A.L., Redford, K.H. & Eisenberg, J.F. (2007) Order Cingulata. In: Gardner, A.L. (Ed.), Mammals of South America. Vol. 1. Marsupial, Xenarthrans, Shrews, and Bats. The University of Chicago Press, Chicago, pp. 128–157.

Wible, J.R. (2010) Petrosal anatomy of the nine-banded armadillo, Dasypus novemcinctus Linnaeus, 1758 (Mammalia, Xenarthra, Dasypodidae). Annals of the Carnegie Museum, 79 (1), 1–28.

Wilson, D.E. & Reeder, D.M. (2005) Mammal species of the world: a taxonomic and geographic reference. 3rd Edition. Johns Hopkins University Press, Baltimore, 2122 pp.

Yepes, J. (1928) Los “Edentata” argentinos. Sistemática y distribución. Revista de la Universidad de Buenos Aires, Series 2a, 1, 461–515.


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