—
## Denny’s Place in Paleogenomic History
Denisova 11 joins a series of discoveries that have collectively transformed the understanding of archaic human populations and their relationships. Placing her in this sequence helps clarify what was already known before 2018, what her discovery added, and what remains to be learned.
Before 2010, the picture of late archaic human populations in Eurasia was built almost entirely from physical anatomy. Neanderthals were well-documented from physical remains across Europe and western Asia. Their anatomy, tool cultures, and time range were known in detail. But the other archaic human populations—the people who lived east of the Neanderthal range, in Central and East Asia—were poorly understood. Some fossils had been found and described as potentially distinct from Neanderthals, but the evidence was fragmentary and interpretations varied.
The Denisovan discovery in 2010 resolved this ambiguity with unexpected clarity, by showing that the genetic approach could identify a population that the physical fossil approach had failed to characterize. The Denisovans were revealed as genetically distinct from Neanderthals, equally closely related to modern humans, and occupying the eastern range. The discovery also established the existence of archaic-modern interbreeding at a population level: Denisovan DNA in modern human populations proved that the groups had met and reproduced.
The Altai Neanderthal genome, published in 2013, showed Denisovan ancestry in a Neanderthal—evidence for the reverse direction of gene flow, from Denisovans into Neanderthals. This established that the gene flow was bidirectional, or at least that it occurred in the direction from Denisovans to Neanderthals at some point. It also showed that the Altai Neanderthal had some indication of inbreeding in their ancestry—the same data that would later be compared with Denisova 11’s data.