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Redescription of the Argyrolagid Microtragulus Bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) Based on New Remains from Northwestern Argentina

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Redescription of the Argyrolagid Microtragulus Bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) Based on New Remains from Northwestern Argentina Palaeontologia Electronica palaeo-electronica.org Redescription of the argyrolagid Microtragulus bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) based on new remains from Northwestern Argentina M. Judith Babot and Daniel A. García-López ABSTRACT This work is based on new dental, cranial, and postcranial remains of the argyro- lagid Microtragulus bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) and dental pieces of Microtragulus sp. coming from late Pliocene-early Pleistocene lev- els of the Uquía Formation exposed at San Roque, Humahuaca (Jujuy Province, Argentina). They were found in association with amphibians, lizards, birds, rodents, and didelphid marsupials, forming an assemblage probably generated by the trophic activity of owls. Specimens were assigned to M. bolivianus based on the following combination of features: M3 subcircular, with a flexus between the mesiolabial lobe and the metacone, paracone and metacone not differentiated in M4, absence of entof- lexid in m1-2 and shallow entoflexid in m3, proportionally large talonid in m4, with a dis- tinguishable distal flexid. Microtragulus bolivianus, initially recorded in Pliocene sediments exposed at the Bolivian Altiplano, was represented only by a mandibular fragment with one incisor and m3-m4. Based on a much more complete sample we present a detailed dental and postcranial description of the species. Since 1904 the family name Argyrolagidae and the generic names Microtragulus and Argyrolagus have been subject of several nomenclatural changes. A revision of these modifica- tions, mainly those occurring in the last 40 years, is also presented. Furthermore, we analyze some mandibular traits of argyrolagids such as the maxillary canal (retrodental canal), a very odd feature present in all the members of the family, which could be related to the passage of a connection between the inferior alveolar and inferior orbital veins, as in some extant mammals. M. Judith Babot. Fundación Miguel Lillo, Miguel Lillo 251, 4000, San Miguel de Tucumán, Tucumán, Argentina; [email protected] and CONICET-Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina Daniel A. García-López. CONICET-Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina, Facultad de Ciencias Naturales, Miguel Lillo 205, 4000, San Miguel de Tucumán, Tucumán, Argentina; [email protected], and Instituto Superior de Correlación Geológica-INSUGEO (CONICET) Babot, M. Judith and García-López, Daniel A. 2016. Redescription of the argyrolagid Microtragulus bolivianus (Metatheria, Polydolopimorphia, Bonapartheriiformes) based on new remains from Northwestern Argentina. Palaeontologia Electronica 19.2.23A: 1-23 palaeo-electronica.org/content/2016/1481-argyrolagid-from-northwestern-argentina Copyright: © July 2016 Society of Vertebrate Paleontology. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. creativecommons.org/licenses/by/4.0/ BABOT & GARCÍA-LÓPEZ: ARGYROLAGID FROM NORTHWESTERN ARGENTINA Keywords: Argyrolagidae; Bonapartheriiformes; Marsupialia; Uquía Formation; Umala Formation; Marplatan SALMA Submission: 3 August 2015 Acceptance: 2 May 2016 INTRODUCTION Microtragulus is the most widely distributed argyrolagid, including records in central and north- Argyrolagidae is an odd group of small fossil western Argentina and the Bolivian Plateau. The metatherians, which inhabited South America current geographical record indicates a disjunct during the Cenozoic. Its temporal range extends distributional pattern comprising coastal as well as from the late Oligocene (Deseadan South Ameri- highland areas. Microtragulus reigi is the only spe- can Land Mammal Age [SALMA]) to the early cies known from lowland areas (the existence of M. Pleistocene (Marplatan SALMA, Sanandresian argentinus remains questionable; see Discussion) subage). The family is only known from Argentina and M. catamarcensis and M. bolivianus are and Bolivia, although the closely related argyro- restricted to mid and high altitudes (García-López lagoid Khlonia is also present in Chile (Ameghino, 1904; Kraglievich, 1931; Simpson, 1970a, 1970b; and Babot, 2015). The oldest record of the genus, Hoffstetter and Villarroel, 1974; Wolff, 1984; Villar- a mandibular fragment of M. catamarcensis, is roel and Marshall, 1988; Goin et al., 2000, 2010; known from ?early Miocene levels exposed at Carlini et al., 2007). Some of the most remarkable Mendoza Province, Argentina (see Garrido et al. features of the group are the presence of an elon- [2014] and García-López and Babot [2015]), and gated snout, large orbits, very large palatal vacu- the more recent records are known from the late ities and wide incisive foramina, inflated tympanic Pliocene-early Pleistocene (M. bolivianus; Marpla- bullae, procumbent lower incisors, hypsodont and tan SALMA, probably Vorohuean subage; Ortiz et highly modified molars, short, and high mandibular al., 2012) and early Pleistocene (M. reigi; Marpla- body, low and short coronoid process, mandibular tan SALMA, Sanandresian subage; Cione and notch long, and maxillary canal present (retroden- Tonni, 2005; Cione et al., 2015). tal canal sensu Hofftsetter and Villarroel, 1974). The genus was initially defined based on the The postcranial skeleton also shows several pecu- morphology of its lower molars (Simpson, 1970a), liarities such as the shortening of the forelimb, the but additional characters of the upper dentition lengthening of the tibia-fibula and metatarsals, and have been recently described for this taxon the reduced number of elements in the posterior (García-López and Babot, 2015). The dental fea- autopodium, specifically the metatarsals and pha- tures that define Microtragulus are the presence of langes, all these features associated with a bipedal a distinct ectoflexid (labial groove) separating the stance and leaping locomotor habits (Simpson, trigonid from the talonid and absent or vestigial 1970a). entoflexid (lingual groove) in m1, trigonid with a For more than a century, several authors stud- dominant metaconid, lower ectostylid and protoco- ied the taxonomy, anatomy, phylogeny, and paleo- nid pointing mesiolabially and separated from the biology of the group (Ameghino, 1904, 1906; ectostylid by a groove (in m1 and/or m2), reduced Rovereto, 1914; Kraglievich, 1931; Rusconi, 1933, talonid with a distinctive tongue-like hypoconid in 1936; Simpson, 1970a, 1970b; Hoffstetter and Vil- m1-2, well-developed entoconid, and hypoconulid larroel, 1974; Wolff, 1984; Villarroel and Marshall, absent or vestigial, and reduced m4, both in length 1988; Sánchez-Villagra and Kay, 1997; Goin et al., and width, bearing a vestigial talonid separated 2000; Sánchez-Villagra et al., 2000; Sánchez-Villa- from the trigonid by ectoflexid and entoflexid. gra, 2001; Abello et al., 2002; Carlini et al., 2007; Regarding the upper dentition, the distinctive fea- Zimicz, 2011; Goin and Abello, 2013). There are tures include a reduced StE in M1, poorly-devel- currently five genera and 12 recognized species. oped paracone coalescent with metacone, reduced Two genera are monospecific (Hondalagus Villar- mesiolabial lobe in M2 and M3, and convex labial roel and Marshall, 1988 and Anargyrolagus Carlini, outline, particularly in M2 (García-López and Pascual, and Goin, 2007) while Proargyrolagus Babot, 2015). Wolff, 1984, Argyrolagus Ameghino, 1904, and Microtragulus bolivianus is the most northern Microtragulus Ameghino, 1904 are represented by species of the genus. It was found by Hoffstetter two or more species (see Table 1). and Villarroel (1974) in levels of the Umala Forma- tion exposed at the locality of Viscachani (Bolivia; 2 PALAEO-ELECTRONICA.ORG TABLE 1. List of the genera and species included in the Family Argyrolagidae and their temporal and geographic distri- bution. Abbreviations: AR, Argentina; BO, Bolivia. Species Temporal distribution Geographic distribution References Proargyrolagus bolivianus Wolf, Late Oligocene (Deseadan) Salla-Luribay Basin (BO) Wolf (1984); Sánchez- 1984 Villagra and Kay (1997) Proargyrolagus argentinus Goin Early Miocene Puesto Almendra, Chubut Goin and Abello (2013) and Abello, 2013 (Colhuehuapian) (AR) Anargyrolagus primus Carlini, Early Miocene Lower Chubut River valley, Carlini et al. (2007); Goin and Pascual, and Goin 2007 (Colhuehuapian) Chubut (AR) Abello (2013) Hondalagus altiplanensis Middle Miocene (Laventan) Quebrada Honda (BO) Villarroel and Marshall Villarroel and Marshall, 1988 (1988); Sánchez-Villagra et al. (2000) Microtragulus argentinus ?Early Pliocene Monte Hermoso, Buenos Ameghino (1904); Simpson Ameghino, 1904 (Montehermosan) Aires (AR) (1970a); Tomassini et al. (2013) M. catamarcensis (Kraglievich, ?Early Miocene - Early 25 de Mayo, Mendoza (AR) Kraglievich (1931); Simpson 1931) Pliocene ?Andalhuala, Catamarca (1970a); Garrido et al. (AR) (2014); García-López and Babot (2015) Microtragulus sp. ?Late Miocene North of Tucumán (AR) García-López and Babot (2015) M. reigi Simpson, 1970a Middle Pliocene – Early Punta San Andrés, Buenos Simpson (1970a); Cione and Pleistocene; Chapadmalan - Aires (AR) Tonni (2005) Marplatan M. bolivianus Hoffstetter and ?Early Pliocene - Early Vizcachani (BO) Hoffstetter and Villarroel Villarroel, 1974 Pleistocene San Roque, Jujuy (AR) (1974); Ortiz et al. (2012) Montehermosan - Marplatan Argyrolagus palmeri Ameghino, Early
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