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The “Condylarths” (Archaic Ungulata, Mammalia) from the Early Palaeocene of Tiupampa (Bolivia): Implications on the Origin of the South American Ungulates

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The “Condylarths” (Archaic Ungulata, Mammalia) from the Early Palaeocene of Tiupampa (Bolivia): Implications on the Origin of the South American Ungulates The “condylarths” (archaic Ungulata, Mammalia) from the early Palaeocene of Tiupampa (Bolivia): implications on the origin of the South American ungulates Christian de MUIZON Laboratoire de Paléontologie, UMR 8569 du CNRS, Muséum national d’Histoire naturelle, 8 rue de Buffon, F-75231 Paris cedex 05 (France) [email protected] Richard L. CIFELLI Oklahoma Museum of Natural History and Department of Zoology, University of Oklahoma, 1335 Asp Ave., Norman, Oklahoma 73019 (USA) [email protected] Muizon C. de & Cifelli R. L. 2000. — The “condylarths” (archaic Ungulata, Mammalia) from the early Palaeocene of Tiupampa (Bolivia): implications on the origin of the South American ungulates. Geodiversitas 22 (1) : 47-150. ABSTRACT The oldest known “condylarth” fauna of the Tertiary of South America comes from basal Palaeocene strata of the Santa Lucía Formation at Tiupampa (Bolivia). It includes five genera and seven species: Molinodus suarezi, Tiuclaenus minutus, T. cotasi n. sp., T. robustus n. sp., Pucanodus gag- nieri, Andinodus boliviensis, and Simoclaenus sylvaticus, n. gen., n. sp. This work presents a thorough revision of all the fossils previously described, which are interpreted in the light of recently discovered specimens including materi- als belonging to three new taxa. The sample contains 33 jaws and 43 isolated teeth. The Tiupampa “condylarths” are included in the Kollpaniinae Marshall, Case & Woodburne, 1990 (= Molinodinae Bonaparte, Van Valen & Kramartz, 1993), a subfamily of the Mioclaenidae (Kollpania tiupampina Marshall & Muizon, 1988, the type species of the type genus of the Kollpaniidae Marshall, Case & Woodburne, 1990 is regarded as a junior syn- onym of Tiuclaenus minutus Muizon & Marshall, 1987). Some of the taxa (e.g., Molinodus suarezi, Tiuclaenus cotasi) are well represented and the cheek dentition of the former is almost completely known. However, other genera such as Andinodus and Simoclaenus are still poorly known (a few lower teeth only in the case of Andinodus). Because of their scant representation, their position within the Kollpaniinae still has to be confirmed by the discovery of new specimens, especially upper molars of Andinodus. The Tiupampa “condylarths” are clearly related to the North American Mioclaenidae, although it is not possible to securely relate them to a specific genus. They share with the North American taxa the following derived characters when compared to Protungulatum: bulbous cusps, the apices of which are approxi- mated in occlusal view; protocone enlarged; stylar shelf greatly reduced or GEODIVERSITAS • 2000 • 22 (1) 47 Muizon C. de & Cifelli R. L. lost; styles reduced and tending to be aligned with para- and metacristae; paraconid appressed against metaconid; entoconid and hypoconulid connate to fused, and forming an obliquely oriented posterolingual crest. The major characteristics of the Kollpaniinae are: metaconid well posterior to proto- conid, strongly inflated, and invading the talonid basin posteriorly; loss of the entocristid; hypoconid extremely inflated, occupying at least the medial half of the talonid (generally more); talonid basin reduced to an obliquely oriented groove that is widely opened lingually; posterior slope of the hypoconulid of m3 less convex than in the other mioclaenids. The Kollpaniinae (Tiupampa “condylarths” and Escribania) are also regarded as closely related to the Didolodontidae and Litopterna (especially the Protolipternidae). The key synapomorphy of the three groups is the morphology and development of the paracristid, which is narrower than half the width of the trigonid, deeply arched posteriorly, projected anteriorly, and attaching on the anterior to anterolingual side of the proto- and paraconids. Study of the Tiupampa “condylarths” confirms that the South American Didolodontidae and Litopterna have their origins in North American Mioclaenidae. A parsimony analysis establishes the monophyly of the clade including the North American Mioclaenidae, the Kollpaniinae (South American Mioclaenidae), the KEY WORDS Didolodontidae, and the Litopterna. This clade is formally designated as a “Condylarths”, new order of mammals, the Panameriungulata. Because no close relationships Didolodontidae, Litopterna, based on dental characters were found with the other groups of South Panameriungulata, American ungulates (or so-called ungulates) — Astrapotheria, Pyrotheria, early Palaeocene, Xenungulata, Notoungulata —, it reinforces the unlikelihood of a single ori- Bolivia, taxonomy, gin of the South American ungulates and therefore brings into question the phylogeny. monophyly of the Meridiungulata. RÉSUMÉ Les « condylarthres » (ongulés archaïques, Mammalia) du Paléocène inférieur de Tiupampa (Bolivie) : implications sur l’origine des ongulés sud-américains. La plus ancienne faune de « condylarthres » du Tertiaire sud-américain pro- vient du Paléocène basal de la Formation Santa Lucía à Tiupampa (Bolivie). Elle comprend cinq genres et sept espèces : Molinodus suarezi, Tiuclaenus (T. minutus, T. cotasi n. sp., T. robustus n. sp.), Pucanodus gagnieri, Andinodus boliviensis et Simoclaenus sylvaticus, n. gen., n. sp. Ce travail présente une revue de tous les fossiles précédemment décrits, lesquels sont interprétés à la lumière de nombreux spécimens récemment découverts, incluant trois nou- veaux taxons. L’échantillon contient 33 mâchoires et 43 dents isolées. Les « condylarthres » de Tiupampa sont inclus dans les Kollpaniinae Marshall Case & Woodburne, 1990 (= Molinodinae Bonaparte, Van Valen & Kramartz, 1993), une sous-famille des Mioclaenidae (Kollpania tiupampina Marshall & Muizon, 1988, l’espèce-type de la famille des Kollpaniidae Marshall, Case & Woodburne, 1990, est considérée comme un synonyme plus récent de Tiuclaenus minutus Muizon & Marshall, 1987). Certain des taxons (Molinodus suarezi, Tiuclaenus cotasi) sont bien représentés et les dents jugales de ces derniers sont presque toutes connues. Cependant, certains genres comme Andinodus et Simoclaenus, sont encore mal connus (par quelques dents inférieures seulement pour Andinodus) et, en raison de la pau- vreté de cet échantillon, leur position au sein des Kollpaniinae devra être confirmée par la découverte de nouveaux spécimens, en particulier de dents supérieures d’Andinodus. Les « condylarthres » de Tiupampa sont, de toute 48 GEODIVERSITAS • 2000 • 22 (1) “Condylarths” from the early Palaeocene of Bolivia évidence, apparentés aux Mioclaenidae nord-américains, bien qu’il ne soit pas possible de les relier avec certitude à un genre précis. Ils partagent avec les formes nord-américaines les caractères dérivés suivants, par comparaison avec Protungulatum : cuspides bulbeuses dont les apex sont rapprochés en vue occlusale ; grand protocone ; plate-forme stylaire très réduite ou perdue ; styles réduits et tendant à s’aligner avec les para- et metacristas ; paraconide plaqué contre le métaconide ; entoconide et hypoconulide accolés, voire fusionnés, et formant une crête postéro-linguale oblique. Les principales caractéristiques des Kollpaniinae sont : très gros métaconide renflé, très posté- rieur au protoconide et envahissant partiellement le bassin du talonide posté- rieurement ; perte de l’entocristide ; hypoconide extrêmement gonflé occupant au moins la moitié labiale du bassin du talonide ; bassin du talonide réduit à un sillon oblique largement ouvert lingualement; flanc postérieur de l’hypoconulide de m3 moins gonflé et moins convexe que chez les autre Mioclaenidae. Les Kollpaniinae (« condylarthres » de Tiupampa et Escribania) sont également considérés comme ayant des relations étroites avec les Didolodontidae et les Litopterna (particulièrement les Protolipternidae). La synapomorphie majeure qui réunit les trois groupes réside dans la morpho- logie et le développement de la paracristide. Celle-ci est plus étroite que la moitié de la largeur du trigonide, profondément arquée postérieurement, for- tement saillante antérieurement et prend attache sur les bords antérieurs à antéro-internes des para- et protoconides. L’étude des « condylarthres » de Tiupampa confirme donc l’origine septentrionale, au sein des Mioclaenidae, des Didolodontidae et des Litopterna d’Amérique du Sud. Une analyse de parcimonie conduit à la monophylie probable du clade incluant les Mioclaenidae d’Amérique du Nord, les Kollpaniinae (Mioclaenidae d’Amérique du Sud), les Didolodontidae et les Litopterna. Le clade est for- MOTS CLÉS mellement désigné comme un nouvel ordre de mammifères, les « Condylarthres », Panameriungulata. Étant donné qu’aucune relation étroite fondée sur des Didolodontidae, Litopterna, caractères dentaires n’a pu être mise en évidence avec les autres groupes Panameriungulata, d’ongulés (ou considérés comme tels) sud-américains, — Astrapotheria, Paléocène inférieur, Pyrotheria, Xenungulata, Notoungulata —, cette étude donne du poids à Bolivie, taxonomie, l’invraisemblance d’une origine unique des ongulés sud-américains et met en phylogénie. question la monophylie des Meridiungulata. INTRODUCTION in situ, having their ancestry among North American “condylarths” that migrated to South The origin of the unique Tertiary ungulate fauna America sometime in the Late Cretaceous or of South America has long captivated the atten- early Tertiary (Simpson 1978). Hence, the less tion of palaeontologists and evolutionary biolo- derived of the South American ungulates, which gists (e.g., Patterson & Pascual 1972; Simpson are included among the “condylarths” (see p. 68 1980). Such groups as notoungulates, for understanding of
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