The phylogenetic position of the Yunxian cranium elucidates the origin of Homo longi and the Denisovans

Diverse Middle Pleistocene forms of Homo coexisted in Africa, Europe, and Asia, and it's controversial whether these fossil humans represent different species or clades. The ~1 Ma old Yunxian 2 fossil from China is crucial for understanding the cladogenesis of Homo and the origin of Homo sapiens . Here, we restored and reconstructed the distorted Yunxian 2 cranium using new technology. The results show that this cranium displays mosaic features of plesiomorphy and apomorphy. Geometric-morphometric, phylogenetic and Bayesian tip-dating analyses including the reconstructed Yunxian 2 suggest that it is an early member of the Asian Homo longi clade, which probably includes the Denisovans, and is the sister group to the Homo sapiens clade. Both the H. sapiens and H. longi clades had deep roots extending beyond the Middle Pleistocene, and the basal position of the Yunxian fossil cranium suggests that it represents a population lying close to the last common ancestor of the two clades.