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Early Steps in the Radiation of Notoungulate Mammals in Southern South America: a New Henricosborniid from the Eocene of Patagonia

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Early Steps in the Radiation of Notoungulate Mammals in Southern South America: a New Henricosborniid from the Eocene of Patagonia Early steps in the radiation of notoungulate mammals in southern South America: A new henricosborniid from the Eocene of Patagonia NICOLÁS BAUZÁ, JAVIER N. GELFO, and GUILLERMO M. LÓPEZ Bauzá, N., Gelfo, J.N., and López, G.M. 2019. Early steps in the radiation of notoungulate mammals in southern South America: A new henricosborniid from the Eocene of Patagonia. Acta Palaeontologica Polonica 64 (3): 597–607. Here we describe a new notoungulate mammal from the early Eocene (Itaboraian SALMA) of Chubut, Argentinian Patagonia, from the localities of Las Flores and Las Violetas Farm, represented by fragments of maxilla and isolated teeth. The specimens were found in the Las Flores Formation, Río Chico Group, and assigned to the Henricosborniidae, a primitive family within the order Notoungulata. Orome deepi gen. et. sp. nov. differs from other henricosborniids in a less developed metacone column in the upper molars, a larger metaloph in the third upper molar and a larger mean size. The morphometric analysis does not show any difference between the individuals of the two studied localities. The new addition to the henricosborniid diversity in Patagonia allows to provide a better understanding of the early radiation of notoungulates in South America. Key words: Mammalia, Notoungulata, Paleogene, Eocene, Patagonia, Argentina. Nicolás Bauzá [[email protected]], División Paleontología de Vertebrados, Museo de La Plata. Paseo del Bosque s/n B1900FWA, La Plata, Buenos Aires, Argentina and Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina. Javier N. Gelfo [[email protected]], División Paleontología de Vertebrados, Museo de La Plata. Paseo del Bosque s/n B1900FWA, La Plata, Buenos Aires, Argentina; Consejo Nacional de Investigaciones Científicas y Técni- cas (CONICET), Argentina; Facultad de Ciencias Naturales y Museo de La Plata, Universidad Nacional de La Plata, Buenos Aires, Argentina. Guillermo M. López [[email protected]], División Paleontología de Vertebrados, Museo de La Plata. Paseo del Bosque s/n B1900FWA, La Plata, Buenos Aires, Argentina; Facultad de Ciencias Naturales y Museo de La Plata, Universidad Nacional de La Plata, Buenos Aires, Argentina. Received 6 November 2018, accepted 25 April 2019, available online 29 July 2019. Copyright © 2019 N. Bauzá et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. An isolated molar from the early Paleocene of Santa Lucía Introduction Formation in Tiupampa, Bolivia (Tiupampan SALMA) was Henricosborniidae Ameghino, 1901 is one of the basal doubtfully assigned to the Henricosborniidae (Muizon et al. families of notoungulates, the most diversified order of ex- 1984; Muizon 1991) and its phylogenetic relationships need tinct South American native ungulates (SAnu). By now, to be confirmed. This scarcity of notoungulates in Tiupampa, the species recognized for this family are characterized al- in contrast to a rich and well preserved, faunistic association most exclusively by dental remains, except for some cranial of marsupials, pantodonts, and mioclaenid kollpaniinaes, specimens of Simpsonotus from the Mealla Formation in could be easily explained by a paleoecological bias. Jujuy province, Argentina (Pascual et al. 1978), and tenta- Despite this record, henricosborniids suddenly thrived tive assigned postcranial remains from Paso del Sapo faunal in the early Eocene in different latitudes of South America. assemblage (Tejedor et al. 2009; Lorente 2015). They are There is no certainty about the cause of notoungulates ab- recorded from several localities of early Eocene Itaboraian sence in the Paleocene record of Patagonia. This could be South American Land Mammal Age (SALMA) in Brazil, at explained due to the temporal gap since the early Paleocene São José de Itaboraí, and in Argentina, in several outcrops units or by a later migration of the group to higher latitudes. in Patagonia (e.g., Simpson, 1948; Tejedor et al. 2009), in Usually recognized genera include Henricosbornia, Peri- north west area (Pascual et al. 1978; Babot et al. 2017), and panto stylops, Oth nielmarshia, and Simpsonotus (McKenna the central Andean region (López 2008). and Bell 1997), with Itaborai therium, Paginula, and Aca- Acta Palaeontol. Pol. 64 (3): 597–607, 2019 https://doi.org/10.4202/app.00565.2018 598 ACTA PALAEONTOLOGICA POLONICA 64 (3), 2019 mana being taxa commonly associated with the family Nomenclatural acts.—This published work and the nomen- (López 2008). Currently the original materials that were clatural acts it contains, have been registered in ZooBank: used to create the species Acamana ambiguus are lost (Bond urn:lsid:zoobank.org:pub:5B1AA50E-D2F3-4E52-BE59- and Vucetich 1983). 4D284744D9AA Since Ameghino (1901) the content and relationships of Henricosborniidae have changed. Originally, Ame ghino placed them within the Prosimiae, alongside the Hyo pso- Material and methods dontidae Selenoconus (Ameghino, 1906). The last revision of the henricosborniids made by Simpson (1948) placed We compared the new remains with the holotypes and syn- them, together with Notostylopidae and Arcto stylopidae, types of the other henricosborniids, and materials referred in the suborder Notioprogonia, which was considered as to other families within the order Notoungulata and basal the most primitive group of notoungulates; Simpson also Paleocene Eutheria (see Appendix 1). In the case of Peri- identified Pantostylopidae and Selenoconus as junior syn- pantostylops minutus a cast and photo graphs of complete onyms of Henricosborniidae and Henricosbornia, respec- upper and lower dental series (AMNH 28494) were used for tively. Nowadays, the Arctostylopidae are considered as the morphometric analysis. an independent radiation of mammals (Cifelli et al. 1989; The phylo genetic analyses were made with TNT ver- Missiaen et al. 2012, but see Kondrashov and Lucas 2004), sion 1.5 (Golo boff et. al. 2008). The matrix utilized was and the group Henricosborniidae–Notostylopidae is consid- modified from Billet (2011), in which we added a couple of ered as paraphyletic. Still, the basal position of the henricos- discrete characters and six landmark configurations (see borniids is recovered in recent phylogenetic analyses, but Appendix 1), one for each upper and lower molar, which due to their fragmentary record and the lack of postcranial we treated as continuous characters (Goloboff and Catalano characteristics, only a few genera (i.e., Henricosbornia and 2016). Because our work is more specific than a phylogeny Simpsonotus) are usually used, recovering the family as of the complete order Notoungulata, we excluded a great paraphyletic (Billet 2011) or polyphyletic (Vera 2015). number of taxa from the original matrix, including only hen- In this work we describe a new genus and species of ricosborniids and a few examples of other families, just to henricosborniid from the Las Flores Formation, Rio Chico get a preliminary glimpse of how Henricosborniidae relates Group in outcrops of two Patagonian localities, Las Flores, to other groups of notoungulates. We also chose Zhelestes in the eastern side of Gran Barranca at the south of Colhué temirkazyk as our outgroup, since it is closer to our group Huapi Lake, and Las Violetas Farm, closer to the shore and of interest. Since the new remains are fragmentary and no near the margin of the San Jorge basin (Raigemborn et. complete dental series was recovered the implementation of al. 2010). This new taxon indicates the presence of a new a single landmark configuration for the geometric morpho- morphological type for the family, increasing their diversity metric analysis was impossible. Photographs of the analyzed and our knowledge of the importance of this group in the materials were compiled into .tps files with tpsUtil software evolutionary history of notoungulates. (Rohlf 2004) and the landmarks were digitized with tps- Dig software (Rohlf 2005). For the upper molars, the two- Institutional abbreviations.—AMNH, American Muse um of dimensional coordinates of eight landmarks were digitized Natural History, New York, USA; CCMGE, Cher nyshev’s over the occlusal surface. For the lower molars, a total of Central Museum of Geological Exploration, Saint Petersburg, six two-dimensional coordinate landmarks were digitized. Russia; FMNH, Field Museum of Natural History, Chicago, The landmark configurations were subjected to a Procrustes USA; MACN, Museo Argentino de Ciencias Naturales, analysis (Rohlf and Slice 1990), in which the configurations Bue nos Aires, Argentina; MCZ, Museum of Comparative are scaled to a centroid size of one, transposed and rotated, Zoology, Harvard College, Cambridge, Massachusetts, USA; so that the sum of squared distances between corresponding MLP, Museo de La Plata, Argen tina; MNHN-SCZ, Santa- landmarks is minimal. The geometric morphometric analy- Cruz collection of the Museum National d’Histoire Naturelle, ses were made with MorphoJ software (Klingenberg 2011), Paris, France; ZIN, Zoological Institute, Russian Academy of including a Principal Component Analysis (PCA) to visual- Sciences, Saint Petersburg, Russia. ize the major trends of shape variation and possible patterns of distribution of specimens along each factor, a Canonical Other abbreviations.—CVA, Canonical
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