Subsequent nuclear DNA sequencing confirmed the distinctiveness of the Denisovans and established the outline of their evolutionary history.

Denisova 4 was a molar tooth — enormous by the standards of modern humans, in the size range associated with archaic human species — from an adult Denisovan.

The genetic analysis of this tooth confirmed that its owner was male and belonged to the same Denisovan population as Denisova 3.

Denisova 5 was a toe bone from a Neanderthal female — an individual whose genome was sequenced to very high quality and has been one of the most important references for Neanderthal genomics.

Strikingly, this Neanderthal genome showed some evidence of distant Denisovan ancestry — suggesting that even before Denny, there had been gene flow between Neanderthals and Denisovans in the opposite direction, with Denisovan genes making their way into Neanderthal populations.