To discover interordinal relationships of living and fossil placental mammals and the time of origin of placentals relative to the Cretaceous-Paleogene (K-Pg) boundary, we scored 4541 phenomic characters de novo for 86 fossil and living species. Combining these data with molecular sequences, we obtained a phylogenetic tree that, when calibrated with fossils, shows that crown clade Placentalia and placental orders originated after the K-Pg boundary. Many nodes discovered using molecular data are upheld, but phenomic signals overturn molecular signals to show Sundatheria (Dermoptera + Scandentia) as the sister taxon of Primates, a close link between Proboscidea (elephants) and Sirenia (sea cows), and the monophyly of echolocating Chiroptera (bats). Our tree suggests that Placentalia first split into Xenarthra and Epitheria; extinct New World species are the oldest members of Afrotheria.

We describe here two new specimens of the mammal Deltatheridium pretrituberculare from the Late Cretaceous period of Mongolia. These specimens provide information on tooth replacement in basal therian mammals and on lower jaw and basicranial morphology. Deltatheroidans, known previously from isolated teeth, partial rostra and jaws from the late Cretaceous of Asia and possibly North America, have been identified variously as eutherians, as basal metatherians (the stem-based clade formed by marsupials and their extinct relatives), or as an outgroup to both eutherians and metatherians. Resolution of these conflicting hypotheses and understanding of the early evolution of the therian lineage have been hampered by a sparse fossil record for basal therians. The new evidence supports metatherian affinities for deltatheroidans and allows a comprehensive phylogenetic analysis of basal metatherians and marsupials. The presence of specialized marsupial patterns of tooth replacement and cranial vascularization in Deltatheridium and the basal phylogenetic position of this taxon indicate that these features are characteristic of Metatheria as a whole. Other morphological transformations recognized here secure the previously elusive diagnosis of Metatheria. The new specimens of Deltatheridium illustrate the effectiveness of fairly complete fossil specimens in determining the nature of early evolutionary events.

Estimates of the time of origin for placental mammals from DNA studies span nearly the duration of the Cretaceous period (145 to 65 million years ago), with a maximum of 129 million years ago and a minimum of 78 million years ago. Palaeontologists too are divided on the timing. Some support a deep Cretaceous origin by allying certain middle Cretaceous fossils (97-90 million years old) from Uzbekistan with modern placental lineages, whereas others support the origin of crown group Placentalia near the close of the Cretaceous. This controversy has yet to be addressed by a comprehensive phylogenetic analysis that includes all well-known Cretaceous fossils and a wide sample of morphology among Tertiary and recent placentals. Here we report the discovery of a new well-preserved mammal from the Late Cretaceous of Mongolia and a broad-scale phylogenetic analysis. Our results exclude Cretaceous fossils from Placentalia, place the origin of Placentalia near the Cretaceous/Tertiary (K/T) boundary in Laurasia rather than much earlier within the Cretaceous in the Southern Hemisphere, and place afrotherians and xenarthrans in a nested rather than a basal position within Placentalia.

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The discovery of Triassic (Norian) turtles from the northwest part of Argentina extends the South American record of turtles by 60 million years. Two skeletons, one almost complete, represent a new genus and species of a basal turtle, Palaeochersis talampayensis. This turtle is a member of the family Australochelidae that was recently erected for Australochelys africanus from the Lower Jurassic of South Africa. Here, it is proposed that Australochelidae is the sister group of Proterochersis plus Casichelydia, that turtles were diverse by the Late Triassic, and that Casichelydia probably originated during the Jurassic.

We describe several fossils referable to Gomphos elkema from deposits close to the Paleocene-Eocene boundary at Tsagan Khushu, Mongolia. Gomphos shares a suite of cranioskeletal characters with extant rabbits, hares, and pikas but retains a primitive dentition and jaw compared to its modern relatives. Phylogenetic analysis supports the position of Gomphos as a stem lagomorph and excludes Cretaceous taxa from the crown radiation of placental mammals. Our results support the hypothesis that rodents and lagomorphs radiated during the Cenozoic and diverged from other placental mammals close to the Cretaceous-Tertiary boundary.

A right petrosal from the ?Aptian or Albian Khoobur locality is referred on the basis of size and morphology to Prokennalestes trofimovi, the earliest eutherian previously known only from dentigerous elements. The petrosal shows a mosaic of primitive and derived features, bearing on the purported therian and eutherian morphotypes. Among the primitive features shared with the Early Cretaceous prototribosphenidan Vincelestes and other more basal taxa that are modified in later eutherians and metatherians are the pattern of basicranial arterial and venous circulation, including a prootic canal and an intrapetrosal inferior petrosal sinus; a vertical paroccipital process; and a fenestra semilunaris, an incomplete wall between the cavum epiptericum and cavum supracochleare. Among the derived features shared with therians is a cochlea coiled through a minimum of 360Њ, with Prokennalestes extending the range of the oldest occurrence of such a coiled cochlea by at least 10 million years. Shared with Late Cretaceous eutherians is a shallow internal acoustic meatus with a thin prefacial commissure. The petrosal referred to Prokennalestes is intermediate in having a reduced anterior lamina and lateral flange, both of which are well developed in Vincelestes and essentially lacking in later eutherians and metatherians. Features previously held to be part of the therian and eutherian morphotypes, such as the absence of the anterior lamina and lateral flange, may have been lost independently in metatherians and in post-Prokennalestes eutherians.