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Type: Article
Published: 2019-08-29
Page range: 401–444
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A new and most complete pampathere (Mammalia, Xenarthra, Cingulata) from the Quaternary of Bahia, Brazil

Programa de Pós-Graduação em Ecologia e Recursos Naturais, Laboratório de Paleoecologia e Paleoicnologia, Departamento de Ecologia e Biologia Evolutiva (DEBE), Universidade Federal de São Carlos (UFSCar), Rodovia Washington Luís, KM 235, São Carlos, 13565-905, SP, Brazil.
Laboratorio de Paleontología de Vertebrados, Centro de Investigaciones Científicas y Transferencia de Tecnología a la Producción (CICYTTP–CONICET), Materi y España, 3105 Diamante, Entre Ríos, Argentina.
Laboratorio de Morfología Evolutiva y Desarrollo (MORPHOS), Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, Buenos Aires, Argentina. División Paleontología de Vertebrados, Museo de La Plata, Paseo del Bosque S/Nº, La Plata, Buenos Aires, Argentina
1Programa de Pós-Graduação em Ecologia e Recursos Naturais, Laboratório de Paleoecologia e Paleoicnologia, Departamento de Ecologia e Biologia Evolutiva (DEBE), Universidade Federal de São Carlos (UFSCar), Rodovia Washington Luís, KM 235, São Carlos, 13565-905, SP, Brazil.
Mammalia armadillo osteology fossil axial skeleton cave


Pampatheriidae is a group of South American native cingulates recorded from the Middle Miocene to the Early Holocene. These animals arrived in North America during the Great American Biotic Interchange. During the Quaternary, at least three genera existed: Tonnicinctus Góis, González Ruiz, Scillato-Yané and Soibelzon, Pampatherium Gervais and Ameghino, and Holmesina Simpson. They are differentiated mainly by craniodental and osteodermal characters. In this paper, we describe a new species of Holmesina from Bahia state, Brazil. Two well-preserved specimens possessing osteoderms associated with their skeletons enabled us to determine the genus and species more reliably. Holmesina cryptae sp. nov. differs from the other Quaternary pampatheres by having the simplest ornamental pattern of osteoderms, sharing characteristics between Pampatherium and Holmesina osteoderms and the most robust skull among the Holmesina species, however preserving the main synapomorphies of the genus. Moreover, the integrity of the remains of H. cryptae sp. nov. enabled us to describe elements of the hyoid apparatus, the clavicle, the entire vertebral column and the pelvis. It was also possible to measure its total axial length (2.2 m) more reliably, which is smaller than previously estimated for other pampatheres. Holmesina cryptae sp. nov. constitutes the sixth species of the genus, and it is the fourth in South America.



  1. Abrantes, É.A.L. & Bergqvist, L.P. (2006) Proposta filogenética para los Dasypodidae (Mammalia: Cingulata). In: Gallo V.P., Brito M., Silva, H.M.A & Figueiredo, F.J. (Eds.), Paleontologia de Vertebrados. Grandes Temas e Contribuições Científicas, Interciência. pp. 261–274.

    Bergqvist, L.P., Abrantes, É.A.L. & Avilla, L.S. (2004) The Xenarthra (Mammalia) of São José de Itaboraí Basin (upper Paleocene, Itaboraian), Río de Janeiro, Brazil. Geodiversitas, 26, 323–337.

    Bordas, A.F. (1939) Craneometría y región auditiva de “Chlamytherium typum” Ameghino. Physis, 14, 447–460.

    Carlini, A.A. & Scillato-Yané, G.J. (1993) Origin and evolution of the “Glyptodontoids”. Journal of Vertebrate Paleontology, 13 (3), 28A.

    Cartelle, C. & Bohórquez, G.A. (1984) Pampatherium paulacoutoi, uma nova espécie de tatu gigante da Bahia, Brasil (Edentata, Dasypodidae). Revista Brasileira de Zoologia, 2, 229–254.

    Casali, D.M. & Perini, F.A. (2017) The evolution of hyoid apparatus in Xenarthra (Mammalia: Eutheria), Historical Biology, 29 (6), 777–788.

    Castellanos, A. (1927) Breves notas sobre los clamidoterios. Publicación del Centro Estudiantes de Ingeniería de Rosario, 1927, 1–8.

    Castellanos, A. (1937) Anotaciones sobre la línea filogenética de los Clamiterios. Publicaciones del Instituto Fisiografía y Geología de la Universidad Nacional Litoral, 26, 1–47.

    Chen, I.H., Kiang, J.H., Correa, V., Lopez, M.I., Chen, P.Y., McKittrick, J. & Meyers, M.A. (2011) Armadillo armor: Mechanical testing and micro-structural evaluation. Journal of the Mechanical Behavior of Biomedical Materials, 4 (5), 713–722.

    De Esteban-Trivigno, S. (2010) Buscando patrones ecomorfológicos comunes entre ungulados actuales y xenartros fósiles. Ameghiniana, 48, 189–209.

    De Esteban-Trivigno, S. (2011) Ecomorfología de xenartros extintos: análisis de la mandíbula con métodos de morfometría geométrica. Ameghiniana, 48, 381–398.

    De Iuliis, G., Bargo, S.M. & Vizcaíno, S.F. (2000) Variation in skull morphology and mastication in the fossil giant Armadillos Pampatherium spp. and allied genera (Mammalia: Xenarthra: Pampatheriidae), with comments on their systematics and distribution. Journal of Vertebrate Paleontology, 20, 743–754.

    De Iuliis, G. & Edmund, A.G. (2002) Vassallia maxima Castellanos, 1946 (Mammalia: Xenarthra: Pampatheriidae), from Puerta del Corral Quemado (Late Miocene to Early Pliocene), Catamarca Province, Argentina. In: Emry, R.J. (Ed.), Cenozoic Mammals of Land and Sea. Smithsonian Institution Press, Washington, D.C., pp. 49–64.

    Delsuc, F., Catzeflis, F.M., Stanhope, M.J. & Douzery, E.J.P. (2001) The evolution of armadillos, anteaters and sloths depicted by nuclear and mitochondrial phylogenies: implications for the status of the enigmatic fossil Eurotamandua. Proceedings of the Royal Society B Biological Sciences, 268, 1605–1615.

    Delsuc, F., Gibb, G.C., Kuch, M., Billet, G., Hautier, L., Southon, J., Rouillard, J.M., Fernicola, J.C., Vizcaíno, S.F., MacPhee, R.D. & Poinar, H.N. (2016) The phylogenetic affinities of the extinct glyptodonts. Current Biology, 26 (4), R155–R156.

    Desbiez, A.L.J. & Kluyber, D. (2013) The role of giant armadillos (Priodontes maximus) as physical ecosystem engineers. Biotropica, 45, 537–540.

    Edmund, AG. (1987) Evolution of the genus Holmesina (Pampatheriidae, Mammalia) in Florida, with remarks on taxonomy and distribution. Pearce-Sellards Series, 45, 1–20.

    Edmund, AG. (1985) The armor of fossil giant armadillos (Pampatheriidae, Xenarthra, Mammalia). Pearce-Sellards Series, 40, 1–20.

    Edmund, A.G. (1996) A review of Pleistocene giant armadillos (Mammalia, Xenarthra, Pampatheriidae). In. Stewart, K.M. & Seymour, K.L. (Eds.), Palaeoecology and Palaeoenvironments of Late Cenozoic Mammals. University of Toronto Press, Toronto, pp. 300–321.

    Edmund, A.G. & Theodor, J. (1997) A new giant Armadillo. 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, pp. 227–232.

    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, D.C., pp. 51–64.

    Fariña, R.A., Vizcaíno, S.F. & Storch, G. (2003) Xenarthra—strange joints for strange mammals. Senckenbergiana Biologica, 83, 1–2.

    Fernicola, J.C., Rinderknecht, A., Jones, W., Vizcaíno, S.F. & Porpino, K. (2017) A new species of Neoglyptatelus (Mammalia, Xenarthra, Cingulata) from the Late Miocene of Uruguay provides new insights on the evolution of the dorsal armor in cingulates. Ameghiniana, 55 (3), 233–252.

    Ferreira, T.M.P., Mayer, E.L., Ferigolo, J., Pereira, J. & Ribeiro, A.M. (2018). Pampatherium humboldtii (Lund, 1839)(Xenarthra, Cingulata) of the Southern Brazilian Quaternary: cranial anatomy and taxonomic remarks. Revista Brasileira de Paleontologia, 21 (2), 158–174.

    Flower, W.H. (1885) An Introduction to the Osteology of the Mammalia. Macmillan and CO, London, 408 pp.

    Galliari, F.C. & Carlini, A.A. (2014) Ontogenetic criteria to distinguish vertebral types on the debated xenarthran synsacrum. Journal of Morphology, 276, 494–502.

    Galliari, F.C. & Carlini, A.A. (2018) Xenarthran synsacrum morphology and evolution. Journal of Mammalian Evolution, 1–12. [published online]

    Galliari, F.C., Carlini, A.A. & Sánchez-Villagra, M.R. (2010) Evolution of the axial skeleton in armadillos (Mammalia, Dasypodidae). Mammalian Biology, 75, 326–333.

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

    Gaudin, T.J. & Lyon, L.M. (2017) Cranial osteology of the pampathere Holmesina floridanus (Xenarthra: Cingulata; Blancan NALMA), including a description of an isolated petrosal bone. PeerJ, 5, e4022.

    Gaudin, T.J. & Wible, J.R. (2006) The phylogeny of living and extinct 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. University of Chicago Press, Chicago, pp. 153–198.

    Gervais, H. & Ameghino, F. (1880) Les mammifères fossiles de l’Amérique du Sud. Savy-Ignon H, Paris-Buenos Aires, 225 pp.

    Gibb, G.C., Condamine, F.L., Kuch, M., Enk, J., Moraes-Barros, N., Superina, M., Poinar, H.N. & Delsuc, F. (2015) Shotgun mitogenomics provides a reference phylogenetic framework and timescale for living xenarthrans. Molecular Biology and Evolution, 33 (3), 621–642.

    Góis, F. (2013) Análisis morfológico y afinidades de los Pampatheriidae (Mammalia, Xenarthra). Ph.D. dissertation, Universidad Nacional de La Plata, La Plata, 312 pp.

    Góis, F., González Ruiz, L.R., Scillato-Yané, G.J. & Soibelzon, E. (2015) A peculiar new Pampatheriidae (Mammalia: Xenarthra: Cingulata) from the Pleistocene of Argentina and comments on Pampatheriidae diversity. PLoS One, 10, 1–25.

    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, 37, 175–186.

    Góis, F., Scillato-Yané, G.J., Carlini, A.A. & Ubilla, M. (2012) Una nueva especie de Holmesina Simpson (Xenarthra, Cingulata, Pampatheriidae) del Pleistoceno de Rondônia, Sudoeste de la Amazonia, Brasil. Revista Brasileira de Paleontologia, 15, 211–227.

    Hill, R.V. (2006) Comparative anatomy and histology of xenarthran osteoderms. Journal of Morphology, 267, 1441–1460.

    Hoffstetter, R. (1956) Contribution à l’étude des Orophodontoidea, Gravigrades cuirassés de la Patagonie. Annales de Paléontologie, 42, 535–636.

    Hoffstetter, R. (1958) Xenarthra. In: Piveteau, J. (Ed.), Traité de Paléontologie. Masson et Cie, Paris, pp. 535–636.

    James, G.T. (1957) An edentate from the Pleistocene of Texas. Journal of Paleontology, 31, 797–808.

    Krmpotic, C.M., Ciancio, M.R., Carlini, A.A., Castro, M.C., Scarano, A.C. & Barbeito, C.G. (2015) Comparative histology and ontogenetic change in the carapace of armadillos (Mammalia: Dasypodidae). Zoomorphology, 134 (4), 601–616.

    McKenna, M.C. & Bell, S.K. (1997) Classification of mammals above the species level. Columbia University Press, New York, 631 pp.

    Mitchell, K.J., Scanferla, A., Soibelzon, E., Bonini, R., Ochoa, J. & Cooper, A. (2016) Ancient DNA from the extinct South American giant glyptodont Doedicurus sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos. Molecular Ecology, 25 (14), 3499–3508.

    Möller-Krull, M., Delsuc, F., Churakov, G., Marker, C., Superina, M., Brosius, J., Douzery, E.J.P. & Schmitz, J. (2007) Retroposed elements and their flanking regions resolve the evolutionary history of xenarthran mammals (armadillos, anteaters, and sloths). Molecular Biology and Evolution, 24, 2573–2582.

    Moura de Jesus, J.F., Igual, E.C., Fernandes, M.A., Dias, W.A.F., Buck, P.V. & Correa, A.J.S. (2015) Estudos tafonômicos em cavernas do Município de Iramaia, Bahia. In: Proceedings of 67th Reunião Anual da Sociedade Brasileira para o Progresso da Ciência, São Carlos.

    Oliveira, É.V. & Bergqvist, L.P. (1998) A new Paleocene armadillo (Mammalia, Dasypodoidea) from the Itaboraí Basin, Brazil. In: Casadío S. (Ed.), Paleógeno de América del Sur y de la Península Antártica, 5, pp. 35–40.

    Paula Couto, C. (1979) Tratado de Paleomastozoologia. Academia Brasileira de Ciências, Rio de Janeiro, 590 pp.

    Paula Couto, C. (1980) Um tatu gigante do pleistoceno de Santa Catarina. Anais da Academia Brasileira de Ciências, 52, 527–531.

    Perea, D. (2005) Pseudoplohohorus absolutus n. sp. (Xenarthra, Glyptodontidae), variabilidad en Sclerocalyptinae y redefinición de una biozona del Mioceno Superior de Uruguay. Ameghiniana, 42, 175–190.

    Rinderknecht, A. (2000) La presencia de osteodermos en las extremidades posteriores de Glyptodon clavipes Owen, 1839 (Mammalia, Cingulata). Ameghiniana, 37, 369–373.

    Robertson, J.S. (1976) Latest Pliocene mammals from Haile XV A, Alachua county, Florida. Bulletin of the Florida State Museum, Biological Sciences, 20, 111–186.

    Rose, K.D. & Emry, R.J. (1993) Relationship of Xenarthra, Pholidota, and fossil “Edentates”: the morphological evidence. In: Szalay, F.S., Novacek, M.J. & McKenna, M.C. (Eds.), Mammal phylogeny: placentals. Springer-Verlag, New York, pp. 81–102.

    Scillato-Yané, G.J. (1982) Los Dasypodidae (Mammalia, Edentata) del Plioceno y Pleistoceno de Argentina. Ph. D. dissertation, Universidad Nacional de La Plata, La Plata, 244 pp.

    Scillato-Yané, G.J. (1986) Los Xenarthra fósiles de Argentina (Mammalia, Edentata). Proceedings of Congreso Argentino de Paleontología y Bioestratigrafía, Mendoza, 4, 151–155.

    Scillato-Yané, G.J. (1976) Sobre un Dasypodidae (Mammalia, Xenarthra) de Edad Riochiquense (Paleoceno superior) de Itaboraí (Brasil). Anais da Academia Brasileira de Ciências, 48, 527–530.

    Scillato-Yané, G.J., Carlini, A.A., Tonni, E.P. & Noriega, J.I. (2005) Paleobiogeography of the late Pleistocene pampatheres of South America. Journal of South American Earth Sciences, 20 (1–2), 132–138.

    Scott, W.B. (1905) Mammalia of the Santa Cruz beds. Paleontology I. Part I Edentata, Part II Insectivora, Part III Glires. Reports of the Princeton University Expeditions to Patagonia 1896–1899. Princeton University Press, Princeton, 489 pp.

    Simpson, G.G. (1930) Holmesina septentrionalis, extinct giant armadillo de Florida. American Museum Novitates, 422, 1–10.

    Soibelzon, E., Zurita, A.E. & Carlini, A.A. (2006) Glyptodon munizi Ameghino (Mammalia, Cingulata, Glyptodontidae): redescripción y anatomía. Ameghiniana, 43, 377–384.

    Tauber, A.A. & Di Ronco, J. (2000) Nuevo hallazgo de placas ventrales de Glyptodon Owen, 1839 (Mammalia, Cingulata, Glyptodontidae) en la provincia de Córdoba, República Argentina. Boletín de la Academia Nacional de Ciencias, 64, 336–347.

    Vieira, A.T., Melo, F., Lopes, H.B.V, Campos, J.C.V, Bomfim, L.F.C, Couto, P.A.A. & Benvenutti, S.M.P. (2005) Projeto cadastro de fontes de abastecimento por água subterrânea, Estado da Bahia. Diagnóstico do Município de Iramaia. CPRM/PRODEEM, Salvador, 28 pp.

    Vizcaíno, S.F., De Iuliis, G. & Bargo, S.M. (1998) Skull shape, mastigatory apparatus, and diet of Vassallia and Holmesina (Mammalia: Xenarthra: Pampatheriidae): when anatomy constrains destiny. Journal of Mammalian Evolution, 5, 291–322.

    Vizcaíno, S.F. (2009) The teeth of the ‘‘toothless’’: novelties and key innovations in the evolution of xenarthrans (Mammalia, Xenarthra). Paleobiology, 35, 343–366.

    Winge, H. (1915) Hordfunde og nulevende Gumlere (Edentata) fra Lagoa Santa, Minas Geraes, Brasilien. Med Udsigt over Gumlernes indbyrdes Slaegtskab, E. Museo Lundii, Copenhagen, 321 pp.