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.

Plesiadapiforms are central to studies of the origin and evolution of primates and other euarchontan mammals (tree shrews and flying lemurs). We report results from a comprehensive cladistic analysis using cranial, postcranial, and dental evidence including data from recently discovered Paleocene plesiadapiform skeletons (Ignacius clarkforkensis sp. nov.; Dryomomys szalayi, gen. et sp. nov.), and the most plesiomorphic extant tree shrew, Ptilocercus lowii. Our results, based on the fossil record, unambiguously place plesiadapiforms with Euprimates and indicate that the divergence of Primates (sensu lato) from other euarchontans likely occurred before or just after the Cretaceous/Tertiary boundary (65 Mya), notably later than logistical model and molecular estimates. Anatomical features associated with specialized pedal grasping (including a nail on the hallux) and a petrosal bulla likely evolved in the common ancestor of Plesiadapoidea and Euprimates (Euprimateformes) by 62 Mya in either Asia or North America. Our results are consistent with those from recent molecular analyses that group Dermoptera with Scandentia. We find no evidence to support the hypothesis that any plesiadapiforms were mitten-gliders or closely related to Dermoptera.Euarchonta ͉ phylogeny ͉ Paromomyidae ͉ Micromomyidae ͉ Paleogene T he origin of Primates represents the first clear step in the divergence of humans from the rest of Mammalia, yet our understanding of this important period in evolutionary history remains limited. The systematic relationships of Paleocene-Eocene plesiadapiforms, which have been considered the ancestors of either Euprimates (primates of ''modern aspect'' or crown-clade primates) (1, 2) or of Dermoptera (3, 4) continue to be debated. Clarifying the position of plesiadapiforms is central to understanding the broader relationships among euarchontan mammals (Primates, Scandentia, Dermoptera), and to testing adaptive hypotheses of primate origins (5, 6) by using direct evidence from the fossil record.Plesiadapiforms are among the most diverse and well sampled Paleogene mammal groups, with Ͼ120 species classified into 11 or 12 families from the Paleocene and Eocene of North America, Europe, Asia, and possibly Africa (7,8). The plesiadapiform dental record is extremely diverse, suggesting correlated diversity in diet and behavior; however, comparatively little is known about the cranial or postcranial morphology of plesiadapiforms [see supporting information (SI) Text, Part 1]. Well preserved crania have been documented for only three families: Plesiadapidae, Microsyopidae, and Paromomyidae (1, 9-11). Postcrania are known from a taxonomically limited sample of North American and European plesiadapids (1), from a sample of North American paromomyids and micromomyids (3, 4, 12) the identification and associations of which are still controversial (13,14), from a recently published North American carpolestid skeleton (15, 16), and from a few other isolated and questionably identified elements (7,17,18). Following the sugges...

In this study, the forelimb of 12 species of tupaiids was analyzed functionally and compared to that of other archontan mammals. Several differences that relate to differential substrate use were found in the forelimb morphology of tupaiids. These differences included shape of the scapula, length and orientation of the coracoid process, size of the lesser tuberosity, shape of the capitulum, length of the olecranon process, and shape of the radial head and central fossa. The forelimb of the arboreal Ptilocercus lowii, the only ptilocercine, is better adapted for arboreal locomotion, while that of tupaiines is better adapted for terrestrial (or scansorial) locomotion. While the forelimb of the arboreal Ptilocercus appears to be habitually flexed and exhibits more mobility in its joints, a necessity for movement on uneven, discontinuous arboreal supports, all tupaiines are characterized by more extended forelimbs and less mobility in their joints. These restricted joints limit movements more to the parasagittal plane, which increases the efficiency of locomotion on a more even and continuous surface like the ground. Even the most arboreal tupaiines remain similar to their terrestrial relatives in their forelimb morphology, which probably reflects the terrestrial ancestry of Tupaiinae (but not Tupaiidae). The forelimb of Urogale everetti is unique among tupaiines in that it exhibits adaptations for scratch-digging. Several features of the tupaiid forelimb reflect the arboreal ancestry of Tupaiidae and it is proposed that the ancestral tupaiid was arboreal like Ptilocercus. Also, compared to the forelimb character states of tupaiines, those of Ptilocercus are more similar to those of other archontans and it is proposed that the attributes of the forelimb of Ptilocercus are primitive for the Tupaiidae. Hence, Ptilocercus should be considered in any phylogenetic analysis that includes Scandentia.