Oldest known euarchontan tarsals and affinities of Paleocene Purgatorius to Primates

Significance Purgatorius has been considered a plausible ancestor for primates since it was discovered, but this fossil mammal has been known only from teeth and jaw fragments. We attribute to Purgatorius the first (to our knowledge) nondental remains (ankle bones) which were discovered in the same ∼65-million-year-old deposits as dentitions of this putative primate. This attribution is based mainly on size and unique anatomical specializations known among living euarchontan mammals (primates, treeshrews, colugos) and fossil plesiadapiforms. Results of phylogenetic analyses that incorporate new data from these fossils support Purgatorius as the geologically oldest known primate. These recently discovered tarsals have specialized features for mobility and provide the oldest fossil evidence that suggests arboreality played a key role in earliest primate evolution. Earliest Paleocene Purgatorius often is regarded as the geologically oldest primate, but it has been known only from fossilized dentitions since it was first described half a century ago. The dentition of Purgatorius is more primitive than those of all known living and fossil primates, leading some researchers to suggest that it lies near the ancestry of all other primates; however, others have questioned its affinities to primates or even to placental mammals. Here we report the first (to our knowledge) nondental remains (tarsal bones) attributed to Purgatorius from the same earliest Paleocene deposits that have yielded numerous fossil dentitions of this poorly known mammal. Three independent phylogenetic analyses that incorporate new data from these fossils support primate affinities of Purgatorius among euarchontan mammals (primates, treeshrews, and colugos). Astragali and calcanei attributed to Purgatorius indicate a mobile ankle typical of arboreal euarchontan mammals generally and of Paleocene and Eocene plesiadapiforms specifically and provide the earliest fossil evidence of arboreality in primates and other euarchontan mammals. Postcranial specializations for arboreality in the earliest primates likely played a key role in the evolutionary success of this mammalian radiation in the Paleocene.