But the article says:
"An earlier analysis of ancient DNA in 40,000 and 50,000-year-old Neanderthal bones, respectively from Spain and Italy, suggested that in their European heartland. But the Neanderthals went extinct around 28,000 years ago – long before modern humans in Europe gained a pale skin. Evidently Neanderthals did not pass these useful local adaptations on to modern humans, despite genetic evidence that the two species interbred."
This article only refers to the relationship between the Neanderthals and contemporary humans. There is, of course, more to the story.
From:
Why Am I Denisovan?
When our ancestors first migrated out of Africa around 60,000 years ago, they were not alone. At least two of our hominid cousins had made the same journey—Neanderthals and Denisovans. Neanderthals, the better known of the two species, left Africa about 300,000 years ago and settled in Europe and parts of western Asia. The Denisovans are a much more recent addition to the human family tree. In 2008, paleoanthropologists digging in a cave in southern Siberia unearthed a 40,000-year-old adult tooth and an exquisitely preserved fossilized pinkie bone that had belonged to a young girl who was between five and seven years old when she died.
Recently, scientists successfully extracted nuclear DNA from the pinkie bone and conducted comparison studies with the genomes of modern humans and Neanderthals. Studies show the girl was closely related to Neanderthals, yet distinct enough to merit classification as a new species of archaic humans, which scientists named “Denisovan” after the cave where the pinkie bone was found. The Denisovan genome also suggests the young girl had brown hair, eyes, and skin.
Surprisingly, the scientists found genetic overlap between the Denisovan genome and that of some present-day east Asians, and, in particular, a group of Pacific Islanders living in Papua New Guinea, known as the Melanesians. It appears the Denisovans contributed between 3 to 5 percent of their genetic material to the genomes of Melanesians. Scientists think that the most likely explanation is that Denisovans living in eastern Eurasia interbred with the modern human ancestors of Melanesians. When those humans crossed the ocean to reach Papua New Guinea around 45,000 years ago, they brought their Denisovan DNA over with them.
If this genetic mixing did occur, the fact that Denisovans were discovered in Siberia but contributed to the genomes of modern humans living in Southeast Asia suggests the species ranged widely across Asia, although their low genetic diversity also indicates their numbers were never very high.
According to one theory, Neanderthals, Denisovans, and modern humans are all descended from the ancient human
Homo heidelbergensis. Between 300,000 to 400,000 years ago, an ancestral group of
H. heidelbergensis left Africa and then split shortly after. One branch ventured northwestward into West Asia and Europe and became the Neanderthals. The other branch moved east, becoming Denisovans. By 130,000 years ago,
H. heidelbergensis in Africa had become
Homo sapiens—our ancestors—who did not begin their own exodus from Africa until about 60,000 years ago.
By comparing the genomes of apes, Denisovans, Neanderthals, and modern humans, scientists hope to identify DNA segments unique to the different groups. Early results already suggest modern humans underwent genetic changes involved with brain function and nervous system development, including ones involved in language development, after splitting from Neanderthals and Denisovans. Identifying and understanding these genetic tweaks could help explain why our species survived and thrived while our close relatives died out."
The only exception I will take with this article is more recent research indicates that homo sapiens migrated out of Africa at least 120,000 years ago based on fossils found in Greece, and possibly older, based on fossils found in Asia.