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We describe an incomplete postcranial skeleton of Catopsbaatar catopsaloides from the ?late Campanian red beds of Hermiin Tsav I, in the Gobi Desert, Mongolia. The skeleton is fragmentary and the preservation of bone surface does not permit reconstruction of the musculature. The studied skeleton contains some parts not preserved or incompletely known in other multituberculate genera, such as a long spinous process in a single lumbar vertebra, which together with long transverse processes preserved in Nemegtbaatar, might indicate that at least some multituberculates had jumping ability. The calcaneus of Catopsbaatar is unusual, differing from most other multituberculates (where known) and other mammals by having a short tuber calcanei, with a large proximal anvil−shaped process strongly bent laterally and ventrally, arranged obliquely with respect to the distal margin of the calcaneus, rather than arranged at 90° to it, as in other mammals. This suggests the presence of strong muscles that attached to the tuber calcanei, perhaps further attesting to jumping abilities in Catopsbaatar. We also describe an unfused pelvic girdle and the first extratarsal spur bone (os cornu calcaris) known in multituberculates.
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Mongolian Late Cretaceous multituberculates (except Buginbaatar) form a monophyletic group for which the suborder Djadochtatheria is proposed. Synapomorphies of Djadochtatheria are: large frontals pointed anteriorly and deeply inserted between the nasals. U-shaped fronto-parietal suture, no frontal-maxilla contact, and edge between palatal and lateral walls of premaxilla. Large, rectangular facial surface of the lacrimal exposed on the dorsal side of the cranial roof is present in all djadochtatherians, but may be a plesiomorphic feature. It is also possible that in djadochtatherians the postglenoid part of the braincase is relatively longer than in other multituberculates. Djadochtatherians have an arcuate p4 (secondarily subtrapezoidal in Catopsbaatar) that does not protrude dorsally over the level of the molars (shared with Eucosmodontidae), I3 placed on the palatal part of the premaxilla (shared with the eucosmodontid Stygimys and the cimolomyid Meniscoessus). The small number of cusps on the upper and lower molars and no more than nine ridges on p4 are possibly plesiomorphies for Djadochtatheria. The djadochtatherian Nessovbaatar multicostatus gen. et sp. n., family incertae sedis from the Barun Goyot Formation is proposed. New specimens of the djadochtatherian genera Kryptobaatar, ?Djadochtatherium, and Kamptobaatar are described and revised diagnoses of these taxa and Sloanbaatar are given. A cladistic analysis of Mongolian Late Cretaceous multituberculates (MLCM), using Pee-Wee and NONA programs and employing 43 dental and cranial characters, 11 MLCM taxa, five selected Late Cretaceous or Paleocene multituberculate genera from other regions, and a hypothetical ancestor based on the structure of Plagiaulacoidea, is performed. The Pee-Wee program yielded two equally fit trees that confirm the monophyly of MLCM excluding Buginbaatar. Kryptobaatar, Djadochtatherium, Catopsbaatar, and Tombaatar form a clade, for which the family Djadochtatheriidae is proposed. Chulsanbaataris the sister taxon of this clade. Bulganbaatar and Nemegtbaatar are the sister group of all other djadochtatherians. Kamptobaatar, Sloanbaatar, and Nessovbaatar form a separate clade in the Pee-Wee tree. The NONAprogram yielded thirty equally parsimonious trees and a strict consensus tree with a poor resolution.
The late Campanian djadochtatherioid multituberculate Catopsbaatar catopsaloides was originally known from three skulls from Hermiin Tsav in the Gobi Desert (Mongolia). Three more skulls from Hermiin Tsav are now available, associated with parts of the previously unknown postcranial skeleton, which will be described separately. We describe herein the skull and dentition of C. catopsaloides, based on all available material, housed in PIN, PM, and ZPAL collections. The genera Catopsbaatar, Djadochtatherium, and Kryptobaatar share several characters, unknown in Tombaatar, such as very long postorbital processes directed postero−laterally and downwards, parietal ridges extending from the posterior margins of the postorbital processes postero−medially, and nuchal crests with prominent lateral wings, incurved anteriorly in the middle, so that the skull in dorsal view is shorter in the middle than laterally. Catopsbaatar shares with Djadochtatherium a very high and prominent anterior zygomatic ridge, and presence of the masseteric protuberance, but differs from it and from other djadochtatherioid genera in having the orbit situated more posteriorly, the intermediate zygomatic ridge adhering to the anterior ridge, and a smaller trapezoidal (rather than crescent−shaped) p4 without ridges; it differs from Kryptobaatar and Djadochtatherium in having three upper premolars (P2 being lost) and shares this last character with Tombaatar. Catopsbaataris known not only from Hermiin Tsav, but also from Baruungoyot Formation of Khulsan, represented there by a single m2. We demonstrate that the separation of the masseter superficialis into two parts, the origins of which leave scars on the lateral wall of the zygomatic arch surrounded by zygomatic ridges, occurs in all the multituberculates (beginning with Paulchoffatiidae), and is regarded as a multituberculate autapomorphy.
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