Abstract
A nearly complete skeleton, including most of the thoracic member bones of the sloth Mylodon darwinii, have been found in Upper Pleistocene strata from Anisacate River, Argentina. The thoracic member bones resemble their homologues in Glossotherium robustum, Paramylodon harlani, and Mylodonopsis ibseni in the following traits: (1) the olecranon is mediolaterally compressed; (2) the radius has an acute styloid process; (3) the radial diaphysis medial border is straight for two thirds of its length; (4) the radial shaft medial border forms an angle with the medial border of the styloid process. The radius presents a distinctive, mostly proximally facing articular circumference. The unfused epiphyses and feeble muscle attachment ridges indicate a sub-adult ontogenetic stage. Deviation of the olecranon and weak M. teres major origin and insertion, suggest a low fossorial specialization. The structure of the thoracic limb bones does not support climbing habits, because pronation-related features are reduced and the humeral head is not prominent. The structure of the radius suggests graviportal adaptations: the proximal head is mediolaterally expanded and the diaphysis straight. A phylogenetic analysis adding thoracic member characters recovers M. darwinii as part of a clade that includes Glossotherium robustum and Paramylodon harlani, but excludes Lestodon armatus. This contrasts with the results of previous analyses focusing on the head skeleton, highlighting the relevance of sampling postcranial characters in phylogenetic analyses of mylodontine sloths.
Kurzfassung
Ein nahezu komplettes Skelett des Faultiers Mylodon darwinii, inklusive der meisten Brust-Extremitätenknochen, wurde in oberpleistozänen Schichten des Anisacate-Flusses in Argentinien gefunden. Es erlaubt erstmals eine detaillierte Beschreibung der Schultergliedmaßen und Unterarmknochen. Diese spiegeln die Homologien in Glossotherium robustum, Paramylodon harlani und Mylodonopsis ibseni mit den folgenden Eigenschaften wider: (1) der Ellenbogen ist mediolateral zusammengedrückt; (2) der Radius hat einen spitzen Styloid-Fortsatz; (3) der radiale Diaphysen-Medialrand ist auf 2/3 der Länge gerade; (4) der radiale Schaft-Medialrand bildet einen Winkel mit dem Medialrand des Styloid-Fortsatzes. Der Radius zeigt einen deutlichen hauptsächlich proximal gerichteten Gelenkumfang. Die unverschmolzenen Epiphysen und schwachen Muskelansatzrücken deuten auf ein sub-adultes ontogenetisches Stadium hin. Grazile Unterarmknochen und die Ausbildung eines schwachen großen Rundmuskels weisen auf eine weniger grabende Spezialisierung hin. Die Struktur der Brust-Extremitätenknochen unterstützt keine Kletterhabitate, da pronations-ähnliche Merkmale reduziert sind und der Humeruskopf nicht markant ist. Die Struktur des Radius deutet auf graviportale Anpassungen hin: der proximale Kopf ist mediolateral ausgeweitet und die Diaphyse ist gerade. Eine phylogenetische Analyse mit Merkmalen der Brust-Extremitätenknochen zeigt, dass M. darwinii näher mit Glossotherium robustum und Paramylodon harlani verwandt ist, als die Lestodontini. Dies steht im Gegensatz zu Resultaten früherer Analysen, welche auf den Schädel fokussiert waren und verdeutlicht damit die Relevanz postcranialer Merkmale in phylogenetischen Analysen von mylodontinen Faultieren.
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Acknowledgements
We thank J. Di Ronco for helping A.A.T. with the field work; M. Sosa and A. Montes for preparing the fossil bones; C. Yonahara for photographing the bones; H. G. McDonald, Á. Miño-Boilini, C. Cartelle, D. Brandoni, R. Hulbert, K. MacKenzie, J. Bar, M. Dantas, R. Juárez, and N. Gardner, for supplying us with relevant literature; D. Peters for editing the English style; H. G. McDonald, R. K. McAfee, and F. Pujos for their useful reviews; the Willi Hennig Society for the free availability of the phylogenetic inference software TNT; CONICET for the postdoctoral fellowships to J.A.H. and J.M.K.; and SeCyT-Universidad Nacional de Córdoba for funding this research (Grant 05/I780).
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Appendices
Appendix 1: Sources of comparative data in the literature
Mylodon darwinii Owen, 1839 | Lehmann-Nitsche (1902), Roth (1899), Nordenskjöld (1900), Studer (1905), Kraglievich (1934), Bargo and Deschamps (1996), Pitana (2011), Saint-André et al. (2010), McAfee (2016) |
Glossotherium robustum (Owen, 1842) | Owen (1842), Rautenberg (1906), Reinhardt (1875), Allen (1913), Lull (1915), Stock (1925), Kraglievich (1934), Hoffstetter (1952), Cartelle (1980), McDonald (1987), Saint-André et al. (2010), Pitana (2011), McAfee (2016) |
Glossotherium wegneri (Spillmann, 1931) | |
Paramylodon harlani (Owen, 1839) | Harlan (1843), Leidy (1855), Allen (1913), Lull (1915), Stock (1925), Kraglievich (1934), Hoffstetter (1952), Hirschfeld (1985), McDonald (1987), Cartelle (1992), Saint-André et al. (2010) |
Mylodonopsis ibseni Cartelle, 1991 | |
Pseudoprepotherium confusum Hirschfeld, 1985 | |
Thinobadistes segnis Hay, 1919 | |
Ocnotherium giganteum (Lund, 1839) | Cartelle (1992) |
Simomylodon uccasamamensis Saint-André, Pujos, Cartelle, De Iuliis, Gaudin, McDonald, and Mamani Quispe, 2010 | Saint-André et al. (2010) |
Lestodon armatus Gervais, 1855 | Gervais (1873), Reinhardt (1875), Kraglievich (1934), McDonald (1987), Cartelle (1992), Bargo et al. (2000), Saint-André et al. (2010), Pitana (2011) |
Brievabradys laventensis Villarroel, 2000 | |
Bolivartherium urumaquensis (Linares, 2004) | Carlini et al. (2006b) |
Unnamed Intertropical “Glossotherium aff. G. lettsomi” sensu Cartelle (1992) | |
“Glossotherium aff. G. robustum” specimen LAP-Q 0004 sensu Pitana (2011) | Pitana (2011) |
“Glossotherium aff. G. robustum” specimen MCN-PV 9718 sensu Pitana (2011) | Pitana (2011) |
Catonyx Ameghino, 1891 | |
Scelidotherium Owen, 1839 | Owen (1839), Burmeister (1881), McDonald (1987), Aramayo (1988), Esteban et al. (1992), Cuenca Anaya (1995), Miño-Boilini (2012), Miño-Boilini et al. (2014), Owen (1842), Cartelle et al. (2009) |
Megatherium americanum Cuvier, 1796 | Owen (1858) |
Megalonyx jeffersoni Desmarest, 1822 | |
Mionothropus cartellei De Iuliis, Gaudin, and Vicars, 2011 | De Iuliis et al. (2011) |
Hapalops Ameghino, 1887 |
Appendix 2: Description of characters used in the phylogenetic analysis
These 45 characters were added to our larger matrix formed by combining Gaudin (2004) and Rincón et al. (2015); therefore, their numbering reflects their inclusion in the total character list.
-
312
Supracondylar foramen in humerus: present (0); absent (1) (Owen 1842).
-
313
Calcaneal facets in astragalus: unfused (0); fused (1) (Owen 1842).
-
314
Dermal ossicles: absent (0); present (1) (Ameghino 1881).
-
315
Contact between articular facets for trapezium-metacarpal I and trapezoid in scaphoid: present (0); absent (1) (Hirschfeld 1985).
-
316
Contact between articular facets for trapezoid and magnum in the scaphoid: present (0); absent (1) (Cartelle 1980).
-
317
Articular facet for lunar on scaphoid: subdivided in two parts (0); unique (1) (modified from Hirschfeld 1985).
-
318
Relationship between mediolateral widths of dorsal and palmar borders of radial articular surface in lunar: dorsal border narrower or not much wider (0); dorsal border more than twice palmar border (1) (Hirschfeld 1985).
-
319
Extent of dorsal border of the facet for the unciform relative to extent of border of dorsal surface between facets for radius and magnum in lunar: shorter (0); longer (1) (Cartelle 1980).
-
320
Angle between articular facets for scaphoid and magnum in lunar: lesser to 145° (0); greater to 145° (1) (Hirschfeld 1985).
-
321
Relief of articular surface for ulna in cuneiform in the axis extending from dorsomedial to palmolateral: approximately straight (0); clearly concave (1) (Cartelle 1980).
-
322
Contact between the articular facets for ulna and pisiform in the cuneiform: present (0); absent (1) (Cartelle 1980).
-
323
Distolateral process of cuneiform: poorly developed (0); well developed (1) (Cartelle 1980).
-
324
Relationship of proximodistal length to dorsopalmar depth in cuneiform: lesser (0); greater (1) (modified from Robertson 1976).
-
325
Relationship between proximodistal length and mediolateral width in the cuneiform: lesser than 0.93 (0); larger than 0.93 (1) (modified from Robertson 1976).
-
326
Prominent ridge on proximal region of the dorsal surface of the cuneiform: present (0); absent (1) (Cartelle 1980).
-
327
Proximal facet of trapezium: concave (0); flat (1) (McDonald 1987).
-
328
Contact between trapezium and trapezoid: present (0); absent (1) (McDonald 1987).
-
329
Exposition in dorsal view of articulated manus of contact between metacarpal II and magnum: present (0); absent (1) (Cartelle 1980).
-
330
Proportions of lateral extremity of articular facet for cuneiform in unciform: dorsopalmarly reduced (0); dorsopalmarly expanded (1) (modified from Hirschfeld 1985).
-
331
Mediolateral development of the axial process of trapezium-metacarpal I: low (0); high (1) (Stock 1925).
-
332
Ratio between length of metacarpal II and dorsopalmar depth of narrowest point in the shaft: more than 0.42 (0); less than 0.42 (1) (modified from Cartelle 1980).
-
333
Relationship between articular facets for metacarpal III and magnum in metacarpal II: contacting (0); separated (1) (Saint-André et al. 2010).
-
334
Articular facet for trapezium in metacarpal II: not concave (0); concave (1) (Saint-André et al. 2010).
-
335
Articular facet for the magnum in metacarpal III: single facet (0); subdivided, with separate dorsoabaxial facet (1) (Hirschfeld 1985).
-
336
Dorsal part of articular facet for metacarpal II in metacarpal III: approximately flat (0); markedly convex (1) (Saint-André et al. 2010).
-
337
Ratio between width of body and bone length in metacarpal III: More than 0.37 (0); 0.37 or less (1) (Haro et al. 2016).
-
338
Expansion of the distal carina upon the dorsal surface of metacarpal III: absent or minimal (0); present, well expanded (1) (Stock 1925).
-
339
Orientation of the distal carina of metacarpal III: dorsopalmar (0) oblique from the middle of the palmar border to the dorsoaxial corner of the distal surface (1) (Stock 1925).
-
340
Proximodistal extension of the articular facet for the metacarpal V in the metacarpal IV: relatively extensive (0); quite narrow (1) (Hirschfeld 1985).
-
341
Relief of palmar region of articular surface for metacarpal III in metacarpal IV: convex (0); concave (1) (Hirschfeld 1985).
-
342
Axial offset in distal articular surface of metacarpal IV: well developed (0); reduced or absent (1) (Hirschfeld 1985).
-
343
Relief of carina on distal articular surface of metacarpal IV in the dorsopalmar direction: flat or concave (0); convex (1) (Rautenberg 1906).
-
344
Palmar extension of articular facet for metacarpal V relative to palmar surface at waist of body in metacarpal IV: does not surpass it (0); surpasses it (1) (Saint-André et al. 2010).
-
345
Ratio between maximal dorsopalmar and axioabaxial extents of proximal phalanx of manual digit II: lesser than 1.27 (0); greater than 1.27 (1) (modified from Stock 1925).
-
346
Dorsopalmar keel in proximal surface of intermediate phalanx in manual digit II: not covered with articular surface (0); mostly covered with articular surface (1) (Cartelle 1980).
-
347
Mediolateral compression in olecranon: absent (0); present (1) (Hirschfeld 1985).
-
348
Distal part of lateral surface of ulna: concave (0); convex (1) (Owen 1842).
-
349
Articular circumference proximal exposition: scarce (0); great (1) (Saint-André et al. 2010).
-
350
Bicipital tubercle location in radius: within proximal third (0); exceeding proximal third (1) (Lull 1915).
-
351
Angle in medial border of radius at pronator crest: well marked (0); not marked (1) (Reinhardt 1875).
-
352
Medial border of radius with convexity expanded along distal half of diaphysis: present (0); absent (1) (Reinhardt 1875).
-
353
Angle in medial border of radius at the distal end: not marked (0); well-marked (1) (Rautenberg 1906).
-
354
Styloid process of radius: not acutely pointed (0); acutely pointed (1) (Rautenberg 1906).
-
355
Separation between articular facets for scaphoid and lunar in the radius: ridge (0); groove (1) (Cartelle 1980).
-
356
Relief of articular surface for scaphoid in radius in mediolateral direction: not concave (0); concave (1) (Rautenberg 1906).
Appendix 3: Data matrix
The following data matrix includes only thoracic member and a few other postcranial skeletal characters. It was analyzed after being fused with those of Gaudin (2004) and Rincón et al. (2015), as explained in the main text. Abbreviations—P, polymorphic, both states 0 and 1 present; X, not applicable.
Appendix 4: Apomorphic thoracic limb skeleton character states, other than from the manus, of recovered clades
- Clade 10::
-
Character 348: 0 → 1. Distal part of lateral surface of ulna convex.
- Clade 11::
-
Character 349: 0 → 1. Articular circumference in radius with great proximal exposure.
Character 352: 0 → 1. Medial border of radius nearly straight along distal half of diaphysis (convergence in Megatheria).
Character 353: 0 → 1. Angle well marked in medial border of radius at the distal end.
Character 354: 0 → 1. Styloid process of radius acutely pointed.
- Clade 12::
-
Character 356: 1 → 0. Articular surface for scaphoid in radius flat or convex in mediolateral direction.
- Clade 12 or 13 (uncertain)::
-
Character 347: 0 → 1. Olecranon mediolaterally compressed (convergence in Megatherium).
Character 351: 0 → 1. Angle in medial border of radius at pronator crest not marked.
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Haro, J.A., Tauber, A.A. & Krapovickas, J.M. Thoracic member (pectoral girdle and forelimb) bones of Mylodon darwinii Owen (Xenarthra, Mylodontidae) from the Late Pleistocene of Central Argentina and their phylogenetic implications. PalZ 91, 439–457 (2017). https://doi.org/10.1007/s12542-017-0350-z
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DOI: https://doi.org/10.1007/s12542-017-0350-z