Abstract

Allotheria are an extinct group of mammaliaforms that originally comprised multituberculates, to which ‘haramiyidans’ and gondwanatherians were later added. Phylogenetic relationships of allotherians have remained controversial since the first reports of the allotherian fossils in the 1800s. Here we report a new euharamiyidan based on a skeletal specimen from the Jurassic Daohugou phase of the Yanliao Biota, China. The superb specimen preserves impressions of differentiated hairs. The pes with impressions of toe skin and claw sheath represents the first such evidence in Mesozoic mammaliamorphs. The remarkable tooth morphologies surpass in complexity any Mesozoic mammals previously known. Phylogenetic analyses place the new species within euharamiyidans from Eurasia and support euharamiyidans, multituberculates, and gondwanatherians as a tripartite monophyletic Allotheria within mammals. This clade is supported not only by their similar molar pattern, but also by many shared derived features of the dental system recognized from discoveries during the last two decades. Allotherians and therians represent the two most successful groups of mammals that are characterized by distinct dental systems. They originated at the latest in the Late Triassic and became diversified in the Middle Jurassic, representing two directions of ecomorphological adaptation during early evolution of mammals.