New records of early Paleocene (earliest Torrejonian) plesiadapiforms from northeastern Montana, USA, provide a window into the diversification of stem primates

Plesiadapiforms (putative stem primates) appear in the fossil record shortly after the Cretaceous/Paleogene boundary and subsequently radiated throughout the Paleocene into a taxonomically and ecomorphologically diverse group. The oldest known plesiadapiforms come from early Puercan (the oldest North American Land Mammal ‘age’ [NALMA] of the Cenozoic) deposits in northeastern Montana, and all records of Puercan plesiadapiforms are taxonomically restricted to members of the Purgatoriidae and the enigmatic genus Pandemonium. Plesiadapiform diversity substantially increased in the following Torrejonian NALMA, but the sparse record of faunas between the Puercan and the well-known middle and late Torrejonian has hampered our understanding of this important interval in early primate evolution. Here we report new plesiadapiform dental fossils from early Torrejonian (To1) deposits from the Tullock Member of the Fort Union Formation in northeastern Montana that record several poorly known taxa including members of the Purgatoriidae, Paromomyidae and Pandemonium, and that document the largest and most diverse assemblage of To1 plesiadapiforms known. We describe a new species of the purgatoriid Ursolestes (Ursolestes blissorum, sp. nov.) that represents the largest plesiadapiform known from the early Paleocene and, among other taxa, provides additional evidence that the temporal range of purgatoriids extended into the Torrejonian. Large sample sizes of the oldest known paromomyid, Paromomys farrandi, allowed us to document intraspecific variability and one undescribed tooth locus. Our observations illuminate changes in dental morphology of some taxa that occurred in To1 and may inform the acquisition of certain diagnostic plesiadapiform dental characters. We evaluate plesiadapiform species richness, mean body mass and body-mass disparity through the Paleocene and reveal unrecognized levels of richness in To1 and a general trend of stable body mass and body-mass disparity. Our findings contribute to documented patterns of plesiadapiform provincialism in the early Paleocene and shed light on the early stages of their Torrejonian radiation.