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Article Systematic Revision of Tropical Brazilian Scelidotheriine Sloths (Xenarthra, Mylodontoidea)

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Article Systematic Revision of Tropical Brazilian Scelidotheriine Sloths (Xenarthra, Mylodontoidea) Journal of Vertebrate Paleontology 29(2):555–566, June 2009 # 2009 by the Society of Vertebrate Paleontology ARTICLE SYSTEMATIC REVISION OF TROPICAL BRAZILIAN SCELIDOTHERIINE SLOTHS (XENARTHRA, MYLODONTOIDEA) CA´ STOR CARTELLE,1 GERARDO DE IULIIS,*,2 and RODRIGO LOPES FERREIRA3 1Museu de Cieˆncias Naturais. Pontifı´cia Universidade Cato´ lica (PUC Minas), [email protected]; 2Faculty of Community Services and Health Sciences, George Brown College of Applied Arts and Technology, 200 King Street East, Toronto, Ontario, Canada M5A 1J5 and Department of Ecology and Evolutionary Biology, University of Toronto, 25 Harbord Street, Toronto, Ontario, Canada M5S 3G5, [email protected]; 3Universidade Federal de Lavras UFLA (Minas Gerais), Brazil ABSTRACT—The analysis of recently recovered ground sloth remains from Minas Gerais, Bahia, and Piauı´ (Brazil) results in a major reinterpretation of the Scelidotheriinae and Megalonychidae of tropical Brazil. Among the specimens collected from Lagoa Santa (Minas Gerais) by Lund during the first half of the 19th century are skeletal elements attributable to two scelidotheriine species, as Lund himself determined and Winge corroborated. This is in contrast to the interpretation of these remains as representing a scelidotheriine and the postcranial remains of a peculiar megalony- chid, as most authors, following Hoffstetter, have believed. A new combination is proposed here for one of the species. Another scelidotheriine recently described from Piauı´ by Gue´rin and Faure is a synonym of this species. INTRODUCTION Reinhardt (1875) based Ocnopus laurillardi on a tibia-fibula that Lund (1846) had assigned to S. bucklandi and a tooth Scelidotheriinae were medium-sized mylodontid ground (ZMUC 1130) of a juvenile giant ground sloth (Megatheriidae) sloths. Their abundant remains indicate that they were common that Lund (1842) had identified as “Megatherium laurillardi” and widespread faunal elements of the South American Plio- (= Eremotherium laurillardi). Reinhardt’s mistake with regard Pleistocene. They possess lobate teeth with flattened grinding to the assignment of this molariform has long been recognized. surfaces, as is general of mylodontids (McDonald and De Iuliis, Cartelle and De Iuliis (1995, 2006) formally recongnized this 2008). Main differences from the other main mylodontid clade, molariform as the holotype of E. laurillardi, making E. lauril- the Mylodontinae, are that the skull is narrow and elongated and lardi the valid name of the intertropical Panamerican giant the dentition is subsequently transversely compressed (McDonald ground sloth. We demonstrate below that the tibia-fibula and Perea, 2002; Dantas and Zucon, 2007). belongs to a scelidotheriine. The monophyly of the Scelidotheriinae is robustly supported In analyzing the material that Lund collected in Lagoa Santa by at least 24 unambiguous synapomorphies, of which 6 are and sent to Denmark, Winge (1915) also recognized two sceli- unique to the clade (Gaudin, 2004). However, several aspects of dotheriine species: Catonyx giganteus and Scelidotherium magnum scelidotheriine systematics have not been satisfactorily resolved, (the equivalent of Lund’s S. bucklandi and S. owenii, respectively). such as the generic level taxonomy and the alpha taxonomy of Hoffstetter (1954) drastically revised Winge’s assessment of this remains principally from the Brazilian Pleistocene. Although material. According to this author, the species identified by Winge both these aspects are considered here, the latter is the main (1915) as S. magnum is distinct from this species (which had been focus of the current report. described from Argentina), and should have been identified as The species of Scelidotheriinae discovered by Lund (1839a,b,c) Scelidodon cuvieri (Lund); the latter species is now placed in Cato- in Lagoa Santa (Minas Gerais, Brazil) have a tortuous history. nyx (McDonald, 1987; see below). Hoffstetter (1954) considered They were initially considered as megalonychids and various Winge’s (1915) C. giganteus invalid because it was based on names have been applied to them (see Systematic Paleontology). an erroneous association of remains. According to Hoffstetter Lund (1846) finally recognized that only two scelidotheriine spe- (1954), the skull, mandible, and teeth belonged to the scelido- cies were present: Scelidotherium owenii Lund, 1846 and Sceli- theriine S. cuvieri (which Winge had recognized as S. magnum), dotherium bucklandi (Lund) Lund, 1846. whereas the postcranial skeletal remains belonged to a megalony- Gervais (1874) based Valgipes deformis on a calcaneum, chid. Winge (1915) assigned to C. giganteus the tibia-fibula that MNHN 7384, recovered from caves in Curvelo (Minas Gerais) Reinhardt (1875) had attributed to O. laurillardi (see above). by the Danish fossil collector P. Claussen. Winge (1915) agreed Hoffstetter (1954) thus proposed a new combination, Ocnopus with the scelidotheriine nature of the calcaneum, but Hoffstetter gracilis (Lund) (see Systematic Paleontology). (1954:761) considered it “clairement diffe´rent de ceux de tous les According to Hoffstetter (1954), this species was known only autres gravigrades connus jusqu‘a` pre´sent,” and viewed it as a from the postcranial remains that Winge (1915) had illustrated megalonychid worthy of its own subfamilial status. Paula Couto and two isolated molariforms (ZMUC 2154, 2406) from Lagoa (1979) considered it as incertae sedis. Santa that Hoffstetter noted and assigned to the species. In support of his hypothesis, Hoffstetter (1954) cited the pres- ence of these two molariforms, in addition to the purported *Corresponding author. megalonychid morphology of the postcranial remains. However, 555 556 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 29, NO. 2, 2009 the presence of numerous skeletal remains of O. gracilis from Institutional Abbreviations—MCL, Museu de Cieˆncias Natur- the Lagoa Santa region (= C. giganteus; 26 individuals are repre- ais da Pontifı´cia Universidade Cato´ lica de Minas Gerais, Belo sented in the Lund collection, according to Winge, 1915) sug- Horizonte; MNHN, Muse´um National d’Histoire Naturelle, Par- gests that if this species were a megalonychid, then numerous is; UFMG, Museu de Histo´ ria Natural, Universidade Federal de molariforms with typical megalonychid morphology should also Minas Gerais, Belo Horizonte; ZMUC, Zoologisk Museum Uni- have been recovered. versitat Copenhagen, Copenhagen. Hoffstetter’s revision and opinions have been accepted by all subsequent authors (e.g., among other, Marshall et al., 1984; MATERIALS AND METHODS Cartelle, 1992). The peculiarities of the alleged megalonychid O. gracilis prompted Hoffstetter (1954) to propose a new sub- This study is primarily a taxonomic revision rather than a family Ocnopodinae, and to promote special status for V. defor- cladistic analysis, although there are, clearly, phylogenetic impli- mis, the species described by Gervais (1874). Hoffstetter (1954) cations. It is based almost entirely on the recently recovered noted its overall axe-shaped tuber, reminiscent of that of Mega- material listed below. We have not included any comparative lonyx, and therefore considered it, too, to be megalonychid. data from the literature on C. cuvieri, except for comparative An incomplete skull definitely pertaining to the Megalonychi- measurements in Winge (1915), because the skull and molari- dae was recovered from a limestone cave in Iporanga (Sa˜o Paulo, form specimens from two different species were, for many years, Brazil) during the 1970s. In an unpublished manuscript (CC, pers. erroneously considered conspecific, and such data might thus be obs.), C. de Paula Couto described this specimen and assigned it unreliable. The material for the new combination proposed here to O. gracilis. This was apparently the ‘missing’ skull of this spe- for the species previously recognized as C. giganteus or O. gracilis is cies, confirming Hoffstetter’s view of its megalonychid affinities. listed as referred material below. Material of C. cuvieri (= Winge’s The most recent development in Brazilian scelidotheriine tax- (1915) Scelidotherium magnum = Hoffstetter’s (1954) Scelidodon onomy is the erection of Scelidodon piauiense by Gue´rin and cuvieri) used for comparative purposes includes: MCL 4265, skull Faure (2004) based on a skull, mandible and humerus from and numerous skeletal elements of a single individual; MCL 4254, Piauı´, Brazil. One may surmise from the discussion above that MCL 4259, skulls; MCL 22682 and 22683, posterior halves of skulls; only one intertropical Brazilian scelidotheriine species, S. cuvieri MCL 22681 and 22686, anterior halves of skulls; mandibles; numer- (= C. cuvieri) has been recognized in the literature over the past ous postcranial elements, including humerus, radius, ulna, elements half century, apart from the material described by Gue´rin and of the manus and pes, femur, tibia, and fibula; UFMG without Faure (2004). This species has also had a long, disputed taxo- number, nearly complete postcranial skeleton; and a cast of the nomic history, described by Paula Couto (1973) as a nomencla- partial calcaneum (MNHN 7384; holotype of V. deformis)figured tural nightmare. Indeed, it may have more synonyms (about 40) by Hoffstetter (1954:761). than any other mammal (Cartelle, 1995). It has already been noted that it was initially interpreted as a Megalonychidae by SYSTEMATIC PALEONTOLOGY Lund (1839b,c and 1840a,b) and as a Scelidotheriinae by Owen XENARTHRA Cope, 1889 (1842), Lund
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