TL;DR: A new Jehol eutherian, Ambolestes zhoui, with a nearly complete skeleton that preserves anatomical details that are unknown from contemporaneous mammals, including the ectotympanic and hyoid apparatus is reported, demonstrating that Sinodelphys is a eutherians and that postcranial differences between Sinodelsphys and the Jehol Eutherian Eomaia are variations among the early members of the placental lineage.
Abstract: Molecular estimates of the divergence of placental and marsupial mammals and their broader clades (Eutheria and Metatheria, respectively) fall primarily in the Jurassic period. Supporting these estimates, Juramaia—the oldest purported eutherian—is from the early Late Jurassic (160 million years ago) of northeastern China. Sinodelphys—the oldest purported metatherian—is from the same geographic area but is 35 million years younger, from the Jehol biota. Here we report a new Jehol eutherian, Ambolestes zhoui, with a nearly complete skeleton that preserves anatomical details that are unknown from contemporaneous mammals, including the ectotympanic and hyoid apparatus. This new fossil demonstrates that Sinodelphys is a eutherian, and that postcranial differences between Sinodelphys and the Jehol eutherian Eomaia—previously thought to indicate separate invasions of a scansorial niche by eutherians and metatherians—are instead variations among the early members of the placental lineage. The oldest known metatherians are now not from eastern Asia but are 110 million years old from western North America, which produces a 50-million-year ghost lineage for Metatheria.
TL;DR: In addition, an incomplete upper dentition of Didelphodon coyi from the Scollard Formation extends the range of this species into the Lancian, co−eval with D. vorax and D. padanicus.
Abstract: Previously undescribed specimens of stagodontid marsupials from Late Cretaceous deposits in Alberta, Canada, reveal new information concerning the upper dentition of Eodelphis spp. and the lower dentition of Didelphodon coyi. Addition− ally, an incomplete upper dentition of D. coyi from the Scollard Formation extends the range of this species into the Lancian, co−eval with D. vorax and D. padanicus. Stagodontids are in accord with other North American Late Cretaceous marsupials for which the appropriate parts are known in lacking diastemata between the canines and the molars while pos− sessing well−developed palatal vacuities, implying that these morphologies characterized ancestral marsupials. If so, the diastema between P1 and P2 in the Asian middle Early Cretaceous “metatherian” Sinodelphys szalayi is convergent on that in Cenozoic didelphids, and the absence of palatal vacuities in South American Paleogene and Neogene borhyaenids is derived, representing a paedomorphic truncation of development. Claims that the Asian Late Cretaceous “metatherian” Deltatheridium pretrituberculare had a marsupial−like dental replacement pattern are tautological, deduced from an a pri− ori acceptance of a marsupial model of replacement to the exclusion of other, no less realistic, alternatives. The new speci− mens of Didelphodon coyi demonstrate that upper and lower premolars occluded broadly, implying that the inflated lin− gual lobes characteristic of Didelphodon premolars evolved primarily as a crushing mechanism, not for passive protec− tion of the gums. Recent speculations that stagodontids were aquatic are not based on credible morphologic or taphonomic evidence and are dismissed, as is speculation that the Judithian species of Eodelphis are sexual morphs of a single species. Current knowledge of Didelphodon compels correction of numerous errors concerning its morphology as presented in recent analyses of marsupial relationships.
TL;DR: Diverse metatherian and eutherian tarsal remains from the Late Cretaceous Bissekty Formation, Kyzylkum Desert, Uzbekistan are described, and their functional and taxonomic properties led to a reassessment of polarity hypotheses of therian, met Heatherian, and e Lutherian cruropedal attributes.
Abstract: Diverse metatherian and eutherian tarsal remains from the Late Cretaceous (middle-late Turonian) Bissekty Formation, Kyzylkum Desert, Uzbekistan (ca 90 MYA) are described. Their functional and taxonomic properties, along with those of other tarsal evidence, led to a reassessment of polarity hypotheses of therian, metatherian, and eutherian cruropedal attributes, and the consequences of this for phylogeny of taxa. There are calcaneal remains of several types of marsupials, and a single astragalus that probably belongs to one of these. This represents greater taxonomic diversity than the dental record suggests. Exceptionally large and distally extending peroneal processes, and small and steeply angled calcaneocuboid articulations facing mediodistally, as seen in the Early Cretaceous Sinodelphys and other Cretaceous and Paleogene taxa, attest not only to the metatherian status of these specimens, but also to the retention of many ancestral therian features, even more so than in both the Tiupampa and Itaborai marsupials of the South American Paleocene (both the calcanea and the astragalus suggest therian traits that are decidedly unlike those of symmetrodonts). Calcanea allocated to the deltatherioid species at Bissekty testify unequivocally to their metatherian affinity. The morphology of the best represented sample of eutherian calcanea from Bissekty, presumably of a number of zhelestid species, appears to be more derived than that of the Late Cretaceous/Paleocene Protungulatum in having a much more reduced peroneal process and a calcaneocuboid articulation that faces distally, oriented nearly at a 90° angle to the long axis of the calcaneus. In fact, this distally facing facet, common in later eutherians (except for lineages in the Paleogene record, and various Carnivora), may not be diagnostic of either the protoeutherian, or even of the protoplacentalian, in spite of its presence in Eomaia. Many putatively “basal” lineages have derived characters, hence such outgroups should not be considered the unequivocal repositories of only ancestral character states.