India's Late Cretaceous fossil mammals include the only undisputed pre-Tertiary Gondwanan eutherians, such as Deccanolestes . Recent studies have suggested a relationship between Deccanolestes and African and European Paleocene adapisoriculids, which have been variably identified as stem euarchontans, stem primates, lipotyphlan insectivores, or afrosoricids. Support for a close relationship between Deccanolestes and any of these placental mammal clades would be unique in representing a confirmed Mesozoic record of a placental mammal. However, some paleogeographic reconstructions place India at its peak isolation from all other continents during the latest Cretaceous, complicating reconstructions of the biogeographic history of the placental radiation. Recent fieldwork in India has recovered dozens of better-preserved specimens of Cretaceous eutherians, including several new species. Here, we incorporate these new specimens into an extensive phylogenetic analysis that includes every clade with a previously hypothesized relationship to Deccanolestes . Our results support a robust relationship between Deccanolestes and Paleocene adapisoriculids, but do not support a close affinity between these taxa and any placental clade, demonstrating that Deccanolestes is not a Cretaceous placental mammal and reinforcing the sizeable gap between molecular and fossil divergence time estimates for the placental mammal radiation. Instead, our expanded data push Adapisoriculidae, including Deccanolestes , into a much more basal position than in earlier analyses, strengthening hypotheses that scansoriality and arboreality were prevalent early in eutherian evolution. This comprehensive phylogeny indicates that faunal exchange occurred between India, Africa, and Europe in the Late Cretaceous-Early Paleocene, and suggests a previously unrecognized ∼30 to 45 Myr “ghost lineage” for these Gondwanan eutherians.

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Extant species of the supraordinal mammal clade Euarchonta belong to the orders Primates, Scandentia, or Dermoptera. The fossil record of euarchontans suggests that they underwent their initial radiation during the Paleocene (65-55 million years ago) in North America, Eurasia, and Africa. The time and place of origin is poorly resolved due to lack of definitive fossils of euarchontan stem taxa. We describe a fragmentary humerus and two fragmentary ulnae from the latest Cretaceous of India that bear significantly on this issue. The fossils are tentatively referred to Deccanolestes cf. hislopi due to their small size and the fact that Deccanolestes is the only eutherian dental taxon to have been recovered from the same locality. The new fossils are used to evaluate the existing behavioral hypothesis that Deccanolestes was arboreal, and the competing phylogenetic hypotheses that Deccanolestes is a stem eutherian versus a stem euarchontan. The humerus resembles those of euarchontans in possessing a laterally keeled ulnar trochlea, a distinct zona conoidea, and a spherical capitulum. These features also suggest an arboreal lifestyle. The ulnar morphology is consistent with that of the humerus in reflecting an arboreal/scansorial animal. Detailed quantitative comparisons indicate that, despite morphological correlates to euarchontan-like arboreality, the humerus of Deccanolestes is morphologically intermediate between those of Cretaceous "condylarthran" mammals and definitive Cenozoic euarchontans. Additionally, humeri attributed to adapisoriculids are morphologically intermediate between those of Deccanolestes and definitive euarchontans. If adapisoriculids are euarchontans, as recently proposed, our results suggest that Deccanolestes is more basal. The tentative identification of Deccanolestes as a basal stem euarchontan suggests that (1) Placentalia began to diversify and Euarchonta originated before the Cretaceous-Tertiary boundary and (2) the Indian subcontinent, Eurasia, and Africa are more likely places of origin for Euarchonta than is North America.

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The sedimentary record documenting the northward drift of India (Late Cretaceous to late Early Eocene) has recently provided important clues to the evolution, radiation, and dispersal of mammals. Here, we report a definitive Late Cretaceous (Maastrichtian) archaic ungulate (Kharmerungulatum vanvaleni genus et species nova) from the Deccan volcano-sedimentary sequences exposed near Kisalpuri village in Central India. This find has important implications for the origin and diversification of early ungulates and raises three possible paleobiogeographic scenarios: (i) Archaic ungulates may have been cosmopolitan in distribution. (ii) Kharmerungulatum might be an immigrant from Western Asia. (iii) Archaic ungulates may have originated in India.