REVISION OF THE MIOCENE CAVIOMORPH RODENTS FROM THE RÍO SANTA CRUZ (ARGENTINEAN PATAGONIA)
DOI:
https://doi.org/10.5710/PEAPA.25.09.2019.299Keywords:
Santacrucian, Province of Santa Cruz, Hystricognathi, Taxonomy, Systematics, Biostratigraphy, EvolutionAbstract
Fossil rodents from the Río Santa Cruz (RSC) classic localities (Santa Cruz Formation, Early–Middle Miocene) are known by the works of F. Ameghino and W.B. Scott since the end of the 19th and beginning of the 20th centuries. These caviomorph rodents have not been comprehensively reviewed since then. In this contribution, we studied new fossil specimens from the cliffs of the RSC (Province of Santa Cruz, Argentina) collected with accurate stratigraphic and geographic data during fieldtrips in 2013 and 2014. An increase in the caviomorph taxonomic richness is observed, based upon our taxonomic study of caviomorphs previously recorded in the RSC. Also, these fieldworks recovered for the first time several taxa previously found in other Santacrucian and even older Patagonian localities (Colhuehuapian, Early Miocene). As a general evolutionary pattern, we note an increase of derived euhypsodont taxa (Prolagostomus, Pliolagostomus, Schistomys, and Eocardia) in Segundas Barrancas Blancas (16.47–15.3 Ma). In addition, a taxonomic replacement of Phanomys by Schistomys is noted between Barrancas Blancas (17.21–16.3 Ma) and Segundas Barrancas Blancas, as well as a notably increase in the abundance of the large Perimys onustus in the latter locality. The present study provides a revision of the caviomorph systematics, and intends to be the starting point to understand the diversity (in all its aspects) and the evolution of this group during the Santacrucian, a major event in the South American mammalian history.
References
Alston, E.R. 1876. On the classification of the order Glires. Proceedings of the Zoological Society of London 2: 347–353.
Álvarez, A., Moyers Arévalo, L, and Verzi, D.H. 2017. Diversification patterns and size evolution in caviomorph rodents. Biological Journal of the Linnean Society 121: 907–922.
Álvarez, A., and Pérez, M.E. 2019. Deep changes in masticatory patterns and masseteric musculature configurations accompanied the eco-morphological evolution of cavioid rodents (Hystricognathi, Caviomorpha), Mammalian Biology DOI: https://doi.org/10.1016/j.mambio.2019.03.009
Álvarez, A., Pérez, S.I., and Verzi, D.H. 2011. Ecological and phylogenetic influence on mandible shape variation of South American caviomorph rodents (Rodentia, Hystricomorpha). Biological Journal of the Linnean Society 102: 828–837.
Álvarez, A., Vieytes, E.C., Becerra, F., Olivares, A.I., Echeverría, A.I., Verzi, D.H., and Vassallo, A.I. 2015. Diversity of craniomandibular morphology in caviomorph rodents. An overview of macroevolutionary and functional patterns. In: A.I. Vasallo, and D. Antenucci (Eds.), Biology of Caviomorph Rodents: diversity and evolution. SAREM Series A, Mammalogical Research (Investigaciones Mastozoológicas), p. 199–228.
Antoine, P.-O., Marivaux, L., Croft D.A., Billet, G., Ganerod, M., Gregory Fanjat, C., Rousse, S., and Salas-Gismondi, S. 2012. Middle Eocene rodents from Peruvian Amazonia reveal the pattern and timing of caviomorph origins and biogeography. Proceedings of the Royal Society B279 1319–1326.
Ameghino, F. 1887a. Enumeración sistemática de las especies de mamíferos fósiles coleccionadas por Carlos Ameghino en los terrenos eocenos de la Patagonia austral. Boletín del Museo de La Plata 1: 1–26.
Ameghino, F. 1887b.Observaciones generales sobre el orden de mamíferos extinguidos sudamericanos llamados toxodontes (Toxodontia) y sinopsis de los géneros y especies hasta ahora conocidos. Anales del Museo de La Plata (Entrega especial, 1936): 1–66, figura1.
Ameghino, F. 1889. Contribución al conocimiento de los mamíferos fósiles de la República Argentina. Actas de la Academia Nacional de Ciencias en Córdoba 6: 1–1027.
Ameghino, F. 1891a. Caracteres diagnósticos de cincuenta especies nuevas de mamíferos fósiles argentinos. Revista Argentina de Historia Natural 1: 129–167.
Ameghino, F. 1891b. Nuevos restos de mamíferos fósiles descubiertos por Carlos Ameghino en el Eoceno inferior de la Patagonia austral. Especies nuevas, adiciones y correcciones. Revista Argentina de Historia Natural 1: 289–328.
Ameghino, F. 1894. Enumération synoptique des espéces des mammiferes fósiles des formations éocenes de Patagonie. Boletín de la Academia Nacional de Ciencias de Córdoba 13: 259–452.
Ameghino, F. 1902. Première Contribution à la connaissance de la faune mammalogique des couches à Colpodon. Boletín de la Academia Nacional de Ciencias en Córdoba 17: 71–138.
Arnal, M. 2012. [Sistemática, filogenia e historia evolutiva de roedores Octodontoidea (Caviomorpha, Hystricognathi) del Oligoceno tardío–Mioceno medio vinculados al origen de la familia Octodontidae. Tesis doctoral, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, 317 p. Unpublished.].
Arnal, M., Kramarz, A.G., Vucetich, M.G., and Vieytes, C.E. 2014. A new Early Miocene octodontoid rodent (Hystricognathi, Caviomorpha) from Patagonia (Argentina) and a reassessment of the early evolution of Octodontoidea. Journal of Vertebrate Paleontology 34: 397–406.
Arnal, M., and Perez, M.E. 2013. A new acaremyid rodent (Hystricognathi, Octodontoidea) from the Middle Miocene of Patagonia (South America) and considerations on the early evolution of Octodontoidea. Zootaxa 3616: 119–134.
Arnal, M., and Vucetich, M.G. 2015a. Main radiation events in Pan-Octodontoidea (Rodentia, Caviomorpha). Zoological Journal of the Linnean Society 175: 587–606.
Arnal, M., and Vucetich, M.G. 2015b. Revision of the fossil rodent Acaremys Ameghino, 1887 (Hystricognathi, Octodontoidea, Acaremyidae) from the Miocene of Patagonia (Argentina) and the description of a new acaremyid. Historical Biology 27: 42–59.
Arnal, M., Vucetich, M.G., Croft, D.A., Bargo, M.S., Fernicola, J.C., and Vizcaíno, S.F. 2017. Systematic revision and evolutionary history of Acarechimys Patterson in Kraglievich, 1965 (Rodentia, Caviomorpha, Octodontoidea). Ameghiniana 54: 307–330.
Bennett, E.T. 1833. On the Chinchillidae, a family of herbivorous rodentia, and on a new genus referrible to it. F.L.S., Sec. Z.S. Communicated May 14, 1833. Transactions of the Zoological Society of London 1: 35–64.
Bertrand, C.O., Flynn, J., Croft, D., and Wyss, A. 2012. Two new taxa (Caviomorpha, Rodentia) from the Early Oligocene Tinguiririca Fauna (Chile). American Museum Novitates 3750: 1–36.
Bostelmann, J.E., Le Roux, J.P., Vásquez, A., Gutiérrez, N.M., Oyarzun, J.L., Carreño, C., Torres, T., Otero, R., Llanos, A., Fanning, C.M., and Herve, F. 2013. Burdigalian deposits of the Santa Cruz Formation in the Sierra Baguales, Austral (Magallanes) Basin: Age, depositional environment and vertebrate fossils. Andean Geology 40: 458–489.
Bowdich, T.E. 1821. An analysis of the natural classifications of Mammalia for the use of students and travelers. Smith p. 115.
Busker, F., Pérez, M.E., Krause, J.M., and Vucetich, M.G. 2017. First record of Banderomys leanzai Kramarz, 2005 (Rodentia, Caviomorpha) in Chubut Province, Patagonia (Argentina). Revista del Museo Argentino de Ciencias Naturales 19: 121–129.
Candela, A.M., Cassini, G.H., and Nasif, N.L. 2013. Fractal dimension and cheek teeth crown complexity in the giant rodent Eumegamys paranensis. Lethaia 46: 369–377.
Candela, A.M., and Nasif, N.L. 2006. Systematics and biogeographic significance of Drytomomys typicus (Scalabrini in Ameghino, 1889) nov. comb., a Miocene Dinomyidae (Rodentia, Hystricognathi) from northeast of Argentina. Neues Jahrbuch für Geologie und Paläontologie 3: 165–181.
Candela, A.M., and Rasia, L.L. 2012. Tooth morphology of Echimyidae (Rodentia, Caviomorpha): homology assessments, fossils, and evolution. Zoological Journal of the Linnean Society 164: 451–480.
Candela, A.M., Rasia, L.L. and Pérez, M.E. 2012. Paleobiology of Santacrucian caviomorph rodents: a morphofunctional approach. In: S.F. Vizcaíno, R.F. Kay, and M.S. Bargo (Eds.), Early Miocene Paleobiology in Patagonia: High-Latitude Paleocommunities of the Santa Cruz Formation. Cambridge University Press, Cambridge, p. 287–305.
Croft, D.A., Flynn, J.J., and Wyss, A.R. 2004. Notoungulata and Litopterna of the Early Miocene Chucal Fauna, northern Chile. Fieldiana: Geology (New Series) 50: 1–52.
Cuitiño, J.I., Fernicola, J.C., Kohn, M.J., Trayler, R., Naipauer, M., Brago, M.S, Kay, R.F., and Vizcaíno, S.F. 2016. U-Pb geochronology of the Santa Cruz Formation (Early Miocene) at the Río Bote and Río Santa Cruz (southernmost Patagonia, Argentina): implications for the correlation of fossil vertebrate localities. Journal of South American Earth Sciences 70: 198–210.
Dunnum, J.L. 2015. Family Cavidae. In: J.L. Patton, U.F.J. Pardiñas, and G. D’Elía (Eds.), Mammals of South America. Vol. 2, Rodents. University of Chicago Press, p. 690–726.
Fernicola, J.C., Cuitiño, J.I., Vizcaíno, S.F., Bargo, M.S., and Kay, R.F. 2014. Fossil localities of the Santa Cruz Formation (Early Miocene, Patagonia, Argentina) prospected by Carlos Ameghino in 1887 revisited and the location of the Notohippidian. Journal of South American Earth Sciences 52: 94–107.
Fields, R.W. 1957. Hystricomorph rodents from the Late Miocene of Colombia, South America. University of California Publications in Geological Sciences 32: 273–404.
Fischer de Waldheim, G. 1817. Adversaria zoological. Memoires de la Societe Imperiale des Naturalistes de Moscou 5: 357–428.
Flynn, J.J., Charrier, R., Croft, D.A., Gans, Ph.B., Herriott, T.M., Wertheim, J.A., and Wyss, A.R. 2008. Chronologic implications of new Miocene mammals from the Cura-Mallín and Trapa Trapa Formations, Laguna del Laja area, south central Chile. Journal of South American Earth Sciences 26: 412–423
Flynn, J.J., Novacek, M.J., Dodson, H.E., Frassinetti, D., McKenna, M.C., Norell, M.A., Sears, K.E., Swisher III, C.C., and Wyss, A.R. 2002. A new fossil assemblage from the southern Chilean Andes: implications for geology, geochronology, and tectonics. Journal of South American Earth Sciences 15: 285–302.
.
Kay, R.F. 2010. A new primate from the early Miocene of Gran Barranca, Chubut Province, Argentina: paleoecological implications. In: R.H. Madden, A.A. Carlini, M.G. Vucetich and R.F. Kay (eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change through the Middle Cenozoic of Patagonia. University of Cambridge Press, Cambridge, UK. p. 220–236.
Kay, R.F., Vizcaíno, S.F., Bargo, M.S., Perry, J.M.G., Prevosti, F., and Fernicola, J.C. 2008. Two new fossil vertebrate localities in the Santa Cruz Formation (late early – early Middle Miocene, Argentina), 51 South latitude. Journal of South American Earth Sciences 25: 87–195.
Kraglievich, L. 1926. Los grandes roedores terciarios de la Argentina y sus relaciones con ciertos géneros pleistocenos de las Antillas. Anales del Museo Nacional de Historia Natural 34: 121–135.
Kraglievich, L. 1940. Los roedores extinguidos del grupo de los Neopiblemidae. Obras de geología y paleontología 3: 739–764.
Kramarz, A.G. 2001. Un nuevo roedor Adelphomyinae (Hystricognathi, Echimyidae) del Mioceno medio–inferior de Patagonia, Argentina. Ameghiniana 38: 163–168.
Kramarz, A.G. 2002. Roedores chinchilloideos (Hystricognathi) de la Formación Pinturas, Mioceno temprano–medio de la provincia de Santa Cruz, Argentina. Revista del Museo Argentino de Ciencias Naturales 4: 167–180.
Kramarz, A.G. 2004. Octodontids and erethizontoids (Rodentia, Hystricognathi) from the Pinturas Formation, Early–Middle Miocene of Patagonia, Argentina. Ameghiniana 41: 199–216.
Kramarz, A.G. 2006a. Eocardiids (Rodentia, Hystricognathi) from the Pinturas Formation, late Early Miocene of Patagonia, Argentina. Journal of Vertebrate Paleontology 26: 770–778.
Kramarz A.G. 2006b. Neoreomys and Scleromys (Rodentia, Hystricognathi) from the Pinturas Formation, late Early Miocene of Patagonia, Argentina. Revista del Museo Argentino de Ciencias Naturales 8: 53–62.
Kramarz, G.A., Vucetich, M.G., and Arnal, M. 2013. A new Early Miocene chinchilloid hystricognath rodent. An approach to the understanding of the early chinchillid dental evolution. Journal of Mammalian Evolution 20: 249–261.
Kramarz, A.G., Vucetich, M.G., Carlini, A.A., Ciancio, M.R., Abello, M.A., Deschamps, C.M., and Gelfo, J.N. 2010. A new mammal fauna at the top of the Gran Barranca sequence and its biochronological significance. In: R.H. Madden, A.A. Carlini, M.G. Vucetich, and R.F. Kay (Eds.), The Paleontology of Gran Barranca, evolution and environmental chang through the middle Cenozoic of Patagonia. Cambridge University, New York, p. 264–277.
Landry, S.O. Jr. 1957. The interrelationships of the new and old world Hystricomorph rodents. University of California Publications in Zoology 56: 1–118.
Loomis, F.B. 1914. The Deseado Formation of Patagonia. Rumford Press, Amherst, 232 p.
López, G.M., Vucetich. M.G., Carlini, A.A., Bond, M., Pérez, M.E., Ciancio, M.R., Pérez, D.J., Arnal, M., and Olivares, A.I. 2011. New Miocene mammal assemblage from Neogene Manantiales Basin, Cordillera Frontal, San Juan, Argentina. In: J.A. Salfity, and R.A. Marquillas (Eds.), Cenozoic Geology of the Central Andes of Argentina. SCS Publisher, Argentina, p. 211–226.
Madden, R.H. 2015. Hypsodonty in Mammals: evolution, geomorphology, and the role of earth surface processes. Cambridge University Press, Cambridge, UK, 423 p.
Madozzo-Jaén, M.C., and Pérez, M.E. 2016. The most ancient caviine rodent (Hystricognathi, Cavioidea) comes from the Late Miocene of Northwest Argentina (South America). Historical Biology DOI: 10.1080/08912963.2016.1166360
Mares, M.A., and Ojeda, R.A. 1982. Patterns of diversity and adaptation in South American Hystricognath rodents. In: M.A. Mares, and H. Genoways (Eds.), Mammalian Biology in South America. University of Pittsburg, Pennsylvania, p. 393–432.
Marshall, L.G. 1976. Fossil localities for Santacrucian (Early Miocene) mammals, Santa Cruz Province, Southern Patagonia, Argentina. Journal of Paleontology 50: 1129–1142.
Marshall, L.G., and Salinas, P. 1990. Vertebrados continentales del Mioceno inferior de Magallanes, Chile. Revista Geológica de Chile 17: 57–87.
Muñoz, N.A., Toledo, N., Candela, A.M., and Vizcaíno, S.F. 2019. Functional morphology of the forelimb of Early Miocene caviomorph rodents from Patagonia. Lethaia 52: 91–106.
Ortiz-Jaureguizar, E., and Cladera, G.A. 2006. Paleoenvironmental evolution of southern South America during the Cenozoic. Journal of Arid Environments 66: 498–532.
Pascual, R., Ortiz-Jaureguizar, E., and Prado, J.L. 1996. Land mammals: paradigm of Cenozoic South American geobiotic evolution. In: G. Arratia (Ed.), Contribution of Southern South America to Vertebrate Paleontology, Müncher Geowissenschaftliche Abhandlungen (A) 30: 265–319. München.
Patterson, B., and Wood, A.E. 1982. Rodents from the Deseadan Oligocene of Bolivia and the Relationships of the Caviomorpha. Bulletin of the Museum of Comparative Zoology 149: 371–543.
Patton, J.L., Pardiñas, U.F.J., and D’Elía, G. 2015. Mammals of South America, vol. 2, Rodents. University of Chicago Press, Chicago, 1336p.
Pérez M.E. 2010a. A new rodent (Cavioidea, Hystricognathi) from the Middle Miocene of Patagonia, mandibular homologies, and the origin of the crown group Cavioidea sensu stricto. Journal of Vertebrate Paleontology 30: 1848–1859.
Pérez M.E. 2010b. [Sistemática, ecología y bioestratigrafía de Eocardiidae (Rodentia, Hystricognathi, Cavioidea) del Mioceno temprano y medio de Patagonia. Tesis Doctoral, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, 385p. Unpublished.].
Pérez, M.E., and Pol, D. 2012. Major Radiations in the Evolution of Caviid Rodents: Reconciling Fossils, Ghost Lineages, and Relaxed Molecular Clocks. PLoS ONE 7 10: e48380.
Pérez, M.E., and Vucetich, M.G. 2012. A revision of the fossil genus Phanomys Ameghino, 1887 (Rodentia, Hystricognathi, Cavioidea) from the early Miocene of Patagonia (Argentina) and the acquisition of euhypsodonty in Cavioidea sensu stricto. Paläontologische Zeitschrift 86: 187–204.
Pérez, M.E., Vucetich, M.G., and Kramarz, A.G. 2010. The first Eocardiidae (Rodentia) in the Colhuehuapian (early Miocene) of Bryn Gwyn (Northern Chubut, Argentina) and the early evolution of the peculiar cavioid rodents. Journal of Vertebrate Paleontology 30: 528–534.
Rasia, L.L. 2016. [Los Chinchillidae (Rodentia, Caviomorpha) fosiles de la Republica Argentina: sistematica, historia evolutiva y biogeografica, significado bioestratigrafico y paleoambiental. Tesis Doctoral, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, 246p. Unpublished.].
Rasia, L.L., and Candela, A.M. 2019. Prolagostomus amplus Ameghino, 1894 is a junior synonym of the Patagonia rodent Pliolagostomus notatus Ameghino, 1887 (Chinchillidae; early Miocene, Santa Cruz Formation). Ameghiniana DOI: 10.5710/AMGH.23.12.2018.3228
Reig, O. 1981. A refreshed unorthodox view of paleobiogeography of South American mammals. Evolution 35: 1032–1035.
Scott, W.B. 1905. Mammalia of the Santa Cruz beds. In: Reports of the Princeton University Expeditions to Patagonia 1896-1899. Part III, Glires 39: 348–487.
Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85: 1–350.
Tauber, A.A. 1997. Los vertebrados de la Formación Santa Cruz (Mioceno inferior–medio) en el extremo sureste de la Patagonia y su significado paleoecológico. Ameghiniana 34: 517–529.
Thomas, O. 1897. On the genera of rodents: an attempt to bring up to date the current arrangement of the Order. Proceedings of the Zoological Society of London (1896) (part IV) 1012 –1028.
Tullberg, T. 1899. Über das System der Nagertiere: eine phylogenetische Studie. Nova Acta Regiae Societatis Scientiarium Upsaliensis 3: 1–514.
Upham, N.S., and Patterson, B.D. 2015. Phylogeny and evolution of caviomorph rodents: a complete timetree for living genera. In: A.I. Vasallo and D. Antenucci (Eds.), Biology of Caviomorph Rodents: diversity and evolution. SAREM Series A, Mammalogical Research (Investigaciones Mastozoológicas), p. 63–120.
Verzi, D.H., Olivares, A.I., and Morgan, C.C. 2014. Phylogeny, evolutionary patterns and timescale of South American octodontoid rodents. The importance of recognizing morphological differentiation in the fossil record. Acta Palaeontologica Polonica 757–769.
Vizcaíno, S.F., Kay, R.F., and Bargo, M.S. 2012. Background for a paleoecological study of the Santa Cruz Formation (late Early Miocene) on the Atlantic coast of Patagonia. In: S.F. Vizcaíno, R.F. Kay, and M.S. Bargo (Eds.), Early Miocene Paleobiology in Patagonia. Cambridge University Press, UK, p. 1–22.
Vucetich, M.G. 1984. Los roedores de la Edad Friasense (Mioceno medio) de Patagonia. Revista del Museo de La Plata (Nueva Serie) 8 Paleontología 50: 47–126.
Vucetich, M.G. l991. Los roedores de Salla y Lacayani (Bolivia) y su correlación con los de otras faunas de edad deseadense (Oligoceno). In: R. Suárez-Soruco (Ed.), Fósiles y Facies de Bolivia, Volume 1, Vertebrados. Santa-Cruz, Bolivia, Revista Técnica de YFPB12, p. 625–629.
Vucetich, M.G., Arnal, M., Deschamps, C.M., Pérez, M.E., and Vieytes, E.C. 2015a. A brief history of caviomorph rodents as told by the fossil record. In: A.I. Vasallo, and D. Antenucci (Eds.), Biology of Caviomorph Rodents: diversity and evolution. SAREM Series A, Mammalogical Research (Investigaciones Mastozoológicas), p. 11–62.
Vucetich, M.G., Dozo, M.T., Arnal, M., and Pérez, M.E. 2014. New rodents (Mammalia) from the Late Oligocene of Cabeza Blanca (Chubut) and the first rodent radiation in Patagonia. Historical Biology: An International Journal of Paleobiology 27: 236–257.
Vucetich, M.G., and Kramarz, A.G. 2003. New Miocene rodents of Patagonia (Argentina) and their bearing in the early radiation of the octodontiform octodontoids. Journal of Vertebrate Paleontology 23: 435–444.
Vucetich, M.G., Kramarz, A.G., and Candela, A.M. 2010. Colhuehuapian rodents from Gran Barranca and other Patagonian localities: the state of the art. In: R.H. Madden, A.A. Carlini, M.G. Vucetich, and R.F. Kay (Eds.), The Paleontology of Gran Barranca: Evolution and Environmental Change through the Middle Cenozoic of Patagonia. University of Cambridge Press, Cambridge, UK, p. 206–219.
Walton, A.H. 1997. Rodents. In: R. Kay, R.H. Madden, R.L. Cifelli, and Flynn, J.J. (Eds.), Vertebrate Paleontology in the Neotropics. The Miocene fauna of La Venta, Colombia. Smithsonian Institution Press, Washington, p. 392–409.
Waterhouse, G.E. 1839. Observations on the Rodentia, with a view to point out the groups, as indicated by the structure of the crania in this order of Mammals. Magazine of Natural History 3: 90–96.
Wood, A.E. 1949. A new Oligocene rodent genus from Patagonia. American Museum Novitates 1435: 1–54.
Wood, A.E. 1955. A revised classification of the rodents. Journal of Mammalogy 36: 165–187.
Wood, A.E., and Patterson, B. 1959. The rodents of the Deseadan Oligocene of Patagonia and the beginnings of South American rodent evolution. Bulletin of the Museum of Comparative Zoology 120: 281–428.
Woods, C.A., and Kilpatrick, C.W. 2005. Infraorder Hystricognathi Brandt, 1855. In: D.E. Wilson, and D.M. Reeder (Eds.), Mammal Species of the World: a Taxonomic and Geographic Reference, 3rd edn. Johns Hopkins University Press, Baltimore, p. 1538–1600.
Zachos, J., Pagani, M., Sloan, L., Thomas, E., and Billups, K. 2001. Trends, rhytms, and aberrations in global climate 65 Ma to Present. Science 292: 686–693.
Downloads
Published
Issue
Section
License
Copyright (c) 2019 Publicación Electrónica de la Asociación Paleontológica Argentina
This work is licensed under a Creative Commons Attribution-NoDerivatives 4.0 International License.
Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a CC Attribution-NonCommercial 4.0 that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.