Thus, it is increasingly looking like there were at least three waves of hominins Out of Africa and into Eurasia, and there could easily have been as many as six (including two modern human waves, two Homo erectus waves, and a wave associated with Homo floresiensis). It isn't inconceivable that new discoveries could support additional waves of hominin Out of Africa migration, although there is really no evidence to support that at this time.
Also, our time resolution degrades as we get into the more remote past. For example, it would be almost impossible to determine if the Homo erectus wave was really a single migration, or was actual several migrations spaced 25,000 to 50,000 years apart from each other. What looks like it might have been two waves of modern humans leaving Africa from our vantage point, would look like a single wave of migration if we were looking at the evidence from a vantage point a million years in the future.
Wave One: Homo Erectus
The second hominin Out of Africa wave occurred around 745,000 to 600,0000 years ago that gave rise to a common Neanderthal-Denisovan lineage (which was probably synonymous with
Homo heidelbergensis) that in turn split into two regional branches. The lineage ancestral to modern humans (who became a distinct species about
300,000 to 200,000 years ago) was not distinct from the lineage ancestral to
Neanderthals and Denisovans at that point.
In this scenario, the Levallois stone tool tradition in Europe was a local Neanderthal development in the Middle Pleistocene, that was apparently not shared with or paralleled by the Denisovan hominins from whom they had separated hundreds of thousands of years before, perhaps because their smaller effective population impeded their ability to improve upon the Acheulian stone tools they used when they left Africa and then split from the Neanderthals.
What About The Denisovan?
The dog that isn't barking in this hypothesis is the Denisovan lineage which appears to have migrated to the east, relative to Neanderthals.
We have extensive fossil and archaeological tool evidence of the Neanderthal branch of the
Homo heidelbergensis tree, and pretty good evidence of the
Homo sapiens branch. But, we have only the slightest evidence of what the Denisovan lineage, which apparently did not see the population expansion that the proto-Neanderthals did, was doing for 400,000 or more years between their separation from Neanderthals and their admixture with modern humans ancestral to Papuan and Australian modern humans.
What was the Denisovan range? How was the Denisovan range impacted by the pre-existing
Homo erectus population? Did the Denisovan people replace
Homo erectus in fairly short order, or did they co-exist with them?
Are
the oldest hominin remains in China that aren't clearly Homo erectus really Denisovans (perhaps highly evolved from the time they left Africa)? Or, are they hybrids of modern humans and Denisovans?
Or, are they some other branch of the hominin tree entirely (e.g. highly evolved
Homo erectus or a previously entirely unknown archaic hominin species from Africa)
, with or without modern human admixture?
This final option puts pressure on the principle of parsimony, since it would require either another wave of Out of Africa hominin migration with no obvious African candidate, or
in situ evolution not demonstrated with transitional fossils anywhere in a 1.5 million year plus time span. But, parsimony has not been a terribly reliable principle for investigators looking at hominin evolution in recent years.
What about Homo floresiensis?
In particular, there are fairly strong indications that
Homo floresiensis may actually have been either a
different more primitive hominin species than
Homo erectus, perhaps a close relation of
Homo habilis who evolved ca. 2,400,000 years ago (about 400,000 years before
Homo erectus, but overlapping with it for about 600,000 years in Africa) and is the oldest definitively identified member of the genus
Homo.
Homo habilis was more adapted for a life in the trees (as opposed to walking longer distances on the ground) than
Homo erectus, used more primitive stone tools than
Homo erectus, and while also an omnivorous meat eater had a diet that was more plant heavy and lighter on meat than
Homo erectus who ate "some tougher foods like leaves, woody plants, and some animal tissues, but that they did not routinely consume or specialize in eating hard foods like brittle nuts or seeds, dried meat, or very hard tubers."
Homo erectus used fire (although it isn't clear just how much mastery they had over it) while
Homo habilis did not - which would have encouraged
Homo habilis to live in places not easily razed by intentionally set forest and brush fires (a key tool used by Australian Aborigines to defeat impressive Australian megafauna).
Homo habilis averaged 3 foot 4 inches to 4 foot 5 inches and averaged about 70 pounds, consistent with
Homo floresiensis which averaged about 3 foot 6 inches. In contrast,
Homo erectus had body proportions similar to modern humans an averaged 4 foot 9 inches to 6 foot 1 inch, and averaged about 88-150 pounds.
So, it is possible to image that
Homo erectus and
Homo habilis could have co-existed in different ecological niches in Asia, with
Homo habilis favoring dense Asian jungles, while
Homo erectus favored lightly wooded area and other more open spaces similar to the African savannah which may have been favored by their ancestors.
Homo floresiensis or its ancestors, could have been a fellow traveler with
Homo erectus or
Homo heidelbergensis, or may have arrived in a separate Out of Africa migration of its own.
Oldowan stone tools, normally associated with
Homo habilis in Africa, are found in Asia from very ancient times until about 200,000 to 300,000 years ago.
The earliest hard evidence of
Homo floresiensis is only about 190,000 years old on the island of Flores (implying that "Hobbits" still left Africa well before the earliest modern human departure from Africa in any scenario), but this could simply be due to a lack of well preserved evidence of them in mainland Asia and island Southeast Asia to the west of the Wallace line.
There is a wide time frame in which the small population of
Homo habilis-like hominins necessary to give rise to the relict population that survived until after modern humans evolved on Flores, could have left Africa (really any time from 2,400,000 years ago to 200,000 years ago).
But, a migration in the roughly 500,000 year long pre-
Homo erectus migration time period would seem to make the most sense. The youngest
Homo habilis remains found in Africa date to 1,400,000 years ago reducing the likelihood of a migration of a relict population long after that date, but after
Homo erectus left Africa,
Homo habilis would face tough competition in Eurasia.
Still, once out of Africa and past the Middle East, there would have been some places in Asia where a small
Homo habilis population could have survived competition with
Homo erectus. Their best strategy may have been to
stay out of the way of the larger members of genus
Homo, such as in jungles better suited to smaller hominins.
Most importantly,
Homo floresiensis could not have been the source of Denisovan ancestry in modern humans in Papua New Guinea and Australia, even though the only location where their remains are observed is perfectly located to fit the distribution of Denisovan ancestry in modern humans.
Some Wild Speculations About A Hobbit/Denisovan/Homo Erectus Pre-History Narrative
Indeed, speculatively, the appearance of
Homo floresiensis on the island of Flores around 190,000 years ago could have been the culmination of their desperate flight from the Denisovans, leading the Hobbits to take a chance on driftwood rafts across the strait to Flores, a trip that the predatory Denisovans weren't willing to risk. Perhaps Denisovans, as they arrived in Southeast Asia were sufficiently smart and versatile to pose a genocidal threat to the Hobbits, even in their jungle-like environments which had protected them in the past because these environments were less well suited to
Homo erectus, who also had less of an edge in intelligence over the Hobbits and were more rigid and ecologically inflexible than the Denisovans.
This scenario also puts Denisovans in the right place, at the right time to explain the geographic pattern of their introgression into modern humans, while making their arrival recent enough and brief enough (assuming that all of the Denisovans were in turn dispatched or assimilated by modern humans) that we would not expect them to leave much of an archaeological trace in Southeast Asia.
This scenario also coincides fairly well with a time period in which there are relatively few traces of
Homo erectus in Asia. About 300,000 to 200,000 years ago, stone tools made with the Oldowan technology, previously associated with Asian
Homo Erectus and with African
Homo habilis disappear. If Denisovans had a lighter archaeological footprint than
Homo erectus, one would expect a gap in the hominin archaeological record during a period in which Denisovans partially or entirely replaced Asia's
Homo erectus population. The last
Homo erectus remains date to
143,000 years ago in Indonesia which would have been the last frontier in a wave of Denisovan expansion if this narrative bears any resemblance to reality.
The
appearance of Acheulian tools in China around 800,000 to 900,000 years ago, could coincide with the arrival of the first Denisovans in Asia, who may have co-existed with
Homo erectus (and perhaps
Homo habilis as well) for about 500,000 years or more in Asia.
The lack of Acheulian tools in Asian
Homo erectus (at least until the Denisovans arrive in Asia)
unlike
Homo erectus in Africa, Europe and South Asia who made this innovation much earlier (and before the migration of
Homo heidelbergensis out of Africa) may have also made it possible for
Homo habilis to avoid the extinction at the hands of
Homo erectus that it experienced in Africa where the adoption of Acheulian tools by
Homo erectus/ergaster and the demise of
Homo habilis come fairly close in time.
A Denisovan surge, after hundreds of thousands of years to the north of
Homo erectus territory and perhaps co-existing with
Homo erectus in China, might reflect some paradigm shifting cultural innovation that the Denisovan's developed around that time in lieu of the Levallois stone tool tradition developed around that time by their Neanderthal cousins in Europe. This unspecified cultural innovation may have given them an edge in hunting and warfare compared to their previous, rather marginal selective fitness in Eurasia that finally made them clearly superior to both
Homo erectus and
Homo habilis who might have been found, until then, in different ecozone of Asia.
Perhaps this Denisovan breakthrough that finally allowed them to become dominant was an innovation with a light archaeological footprint such as something comparable to the development of bamboo weapons, or frog poison blow darts, or an advance in language abilities, as opposed to more advanced stone tools than they had from the outset. Or perhaps, the break through was a matter not of technology, but of a major climate event that the more intelligent Denisovans managed to survive better than the duller and more rigid
Homo erectus and
Homo habilis.
Perhaps, by the time modern humans arrived in Asia, the Denisovan has brought about the extinction of
Homo erectus and
Homo habilis in all but a few small relict communities in Asia, a favor that modern humans returned by causing the extinction and/or assimilation of all of the remaining Denisovans around 75,000 to 65,000 years ago, or perhaps earlier, with an initial boost created by the ecological disruption arising from the Toba super volcano explosion.
A third hominin Out of Africa wave brought
Homo sapiens, with a first
Homo sapien Out of Africa wave about 125,000 to 100,000 years ago and possibly with a second wave of
Homo sapiens to leave Africa about 75,000 to 60,000 years ago. It isn't clear if the second
Homo sapiens Out of Africa wave is derived from the first
Homo sapiens Out of Africa wave, or if the second
Homo sapiens Out of Africa wave was separate from the first, with the first wave being largely a dead end leaving only some Middle Eastern relics and some admixture of ancestors of the Altai Neanderthals. We have very little evidence to guide us one way or the other.
Loose ends in Africa
It is fairly likely that other archaic hominin species with admixed with modern humans within Africa who can only be identified as
"ghost populations" in African whole genomes right now may also have arisen from this
Homo heidelbergensis lineage as well with one study noting that: "On the basis of an excess of shared derived alleles between San, Neanderthal, and Denisova we suggest that a third archaic population related more closely to Neanderthal and Denisova than to modern humans introgressed into the San genomes studied here." The admixture with the San probably took place in the last 100,000 years.
There was also probably another archaic admixture, possibly with a different "ghost species" in Africa, that impacted
the African Pygmy genome.
An unknown archaic species that broke off from the lineage that includes
Homo heidelbergensis about 3,450,000 years ago (quite possibly, given the timing, the pre-
Homo species affectionately known as
"Lucy" a.k.a.
Australopithecus afarensis or
Australopithecus africanus, for whom 3.45mya is a bit of a stretch, but which persists much later than Lucy) may have introgressed into a quite basal version of
Homo heidelbergensis.
Homo naledi in Southern Africa is a recently discovered diminutive species of archaic hominins that still existed when modern human evolved (those found are from
300,000 to 200,000 years ago), although their remains suggest that they are part of a fairly archaic and basal branch of the human evolutionary tree, with estimates based upon how archaic its features were in the range of
2,500,000 to 912,000 years ago.
Homo rudolfensis known from only one decent set of remains from about 1,850,000 years ago that was discovered in Kenya in 1986 seems similar to
Homo habilis but with a larger brain case. It isn't clear exactly where this specimen belongs in the evolutionary tree.
Other Archaic Hominin Stories
In other archaic hominin news, John Hawks muses over
why stable hybrid zones apparently didn't emerge between Neanderthals, Denisovans and modern humans respectively.
As the above discussions suggest, however, even if stable hybrid zones didn't emerge in Eurasia, they probably did exist for some extended periods of time in parts of Africa, where hominins originated and hominin populations were larger. Some of the populations in Eurasia may have been too greatly diverged from each other, and too small in effective population size themselves, to give rise to reasonably sized stable hybrid populations.