Cracks in the Out of Africa Theory?

The most exciting finding of the past two decades in biological anthropology has been, without a doubt, evidence for interbreeding between Homo sapiens, Neanderthals, Denisovans, etc. and the sheer multiplicity of new hominin species being uncovered.

Exactly where the lines between species lie is a bit of a matter of semantic debate–where exactly did our ancestors end and Homo sapiens begin? Should we classify Neanderthals and Sapiens as one species if we interbred? etc–but if we accept the current classifications as decent approximations, we have:

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Source: Wikipedia

Sorry, I realized it would be much more efficient if I just grabbed the family tree off Wikipedia instead of copying it over bit by bit. The top part of the tree got cut off, so I’ll note that Homini (6.3 million years ago) includes us + chimps, while Hominina (5.7 million years ago) has no chimps, but includes australopithecines. Gorillas are way back in Homininae, with an e. Homo, our genus, includes all of the “human” species, but usually doesn’t include australopithecines.

There is further debate on exactly who descended from whom. We’re finding new fossils all the time, which is quite exciting, but our current record is not nearly as complete as we’d like it to be. So sometimes branches get moved around or re-categorized as more data comes to light.

The most recent notable additions to our genus are the Denisovans, Homo floresiensis, Homo luzonensis, Homo Naledi, Homo doesn’t have a name yet, and more Denisovans.

You have probably heard of the “hobbit,” Homo floresiensis. The remains we have uncovered of this diminutive hominin are remarkably good, including a skull in great condition (despite some damage caused after excavation.) They lived on the island of Flores from about 200,000 to 50,000 years ago (though their arrival may get pushed back considerably because there are stone tools on Flores that are much older–700,000 years old–we just don’t know yet who used them.)

The hobbits are remarkable in multiple ways. First, they lived in an area that was not connected to the mainland by any landbridges–that is, they had to swim, boat, or otherwise be carried to their island. I am skeptical of the idea of anyone surviving a tsunami as a means of populating an island, but they arrived in an era when, as far as we know, humans had yet to build boats. So perhaps their ancestors were among the first humans to build boats, and we just haven’t found the remains of their crafts (wood being a material that degenerates very quickly.)

Second, the Hobbits are most likely descended from Homo erectus, who lived nearby on mainland Indonesia (at the time, connected to the rest of Asia via a landbridge), but are morphologically very different. They are tiny–shorter than pygmies, Homo erectus, or even australopithecines.

There is much debate about whether they are descended directly from erectus, or part of a sister-clade to erectus that descended from a common ancestor. It was previously believed that erectus was the first hominin to leave Africa, but if Floresiensis is not descended from erectus, Flores could be the first.

Now a similarly diminutive hominin has turned up in the Philippines, also past a significant water barrier that would require some effort to cross. It has been dubbed Homo luzonensis. Not much is known, yet, about luzonensis, (we haven’t found as many of its bones), but what we do know is tantalizing:

It was soon apparent to Détroit that the remains featured a puzzling mosaic of traits both modern and ancient. “Each of the features [of Homo luzonensis] corresponds to some hominin or another,” he says. “But the combination makes for something really unique. There’s no known species with this same suite of features.”

They’re small, possibly even smaller than the Hobbits. Their feet resemble australopithecines, but australopithecines supposedly died out a couple million years before Luzonensis arrived on the scene. And their teeth were “remarkably uniform,” which probably sounds boring to anyone who isn’t a dentist, but provides strong evidence of them being a different species.

Two island dwelling species in the same area supports the idea that their ancestors either developed boats or were remarkably skilled at surviving tsunamis, and that southeast Asia was a remarkable hotspot of hominin diversity.

And then there are the Denisovans!

Denisovans are mysterious because we have so few of their bones–a chunk of skull was recently uncovered, but we have no jaws, no faces, no ribs, etc–so we don’t have a good idea of what they looked like. What we do have are Denisovan DNA (extracted from those fragments of skeletons) and traces of Denisovan DNA in modern humans.

Oddly, those Denisovan bones turned up in Siberia (a good place for preserving old bones, but not such a great place for humans adapted to warm climates) while the humans with Denisovan DNA live in modern Papua New Guinea and nearby areas.

The obvious answer to this puzzle is that both the Denisovans had a much broader range than one cave in Siberia and the ancestors of modern folks from PNG used to live in different areas than they do now.

A more detailed analysis of PNG DNA was recently released, which reveals three separate, significant groups of Denisovans who interbred with sapiens:

… modern Papuans carry hundreds of gene variants from two deeply divergent Denisovan lineages that separated [from each other] over 350 thousand years ago. Spatial and temporal structure among these lineages suggest that introgression from one of these Denisovan groups predominantly took place east of the Wallace line and continued until near the end of the Pleistocene. A third Denisovan lineage occurs in modern East Asians. This regional mosaic suggests considerable complexity in archaic contact, with modern humans interbreeding with multiple Denisovan groups that were geographically isolated from each other over deep evolutionary time.

The Wallace line is a place that’s too deep for a landbridge, and thus the area to the east was an island even during the Ice Age. In other words, it looks like Denisovans could use boats. (Or survive tsunamis, pfft.)

Next, we sought to retrieve dates of divergence between D1, D2, and the Altai Denisovan genome … to encompass two deeply divergent Denisovan-related components, our best fitting model indicates that D1 and D2 split from the Altai Denisovan approximately 283 kya … respectively (Figure 4B). While clearly branching off the Denisovan line, D2 diverged so closely to the Neanderthal-Denisovan split that it is perhaps better considered as a third sister group… For context, even the youngest of these divergence times is similar to the evolutionary age of anatomically modern humans … Our model implies substantial reproductive separation of multiple Denisovan-like populations over a period of hundreds of thousands of years. … 

The genetic diversity within the Denisovan clade is consistent with their deep divergence and separation into at least three geographically disparate branches, with one contributing an introgression signal in Oceania and to a lesser extent across Asia (D2), another apparently restricted to New Guinea and nearby islands (D1), and a third in East Asia and Siberia (D0). This suggests that Denisovans were capable of crossing major geographical barriers, including the persistent sea lanes that separated Asia from Wallacea and New Guinea. They therefore spanned an incredible diversity of environments, from temperate continental steppes to tropical equatorial islands.

(We will probably reclassify some of the older fossils from Asia as Denisovans once we figure out what they looked like.)

Then we have Homo naledi, from South Africa. Naledi lived around 250,000 years ago, about the same time as Homo sapiens were differentiating from their ancestors. We have a wonderful array of Homo naledi fossils, preserved in the bottom of a cave pit. If they were placed here intentionally, this was pretty advanced behavior, though I wonder if perhaps they just got lost in the cave from time to time and then died in the pit.

At about 5 ft tall, (male height) naledi was short, but not nearly as short as Floresiensis, and taller than some groups of sapiens. Its skull was significantly smaller than a modern skull, however, and I find it odd that, out of the thousands of bones and fragments discovered, we have not yet recovered much of the front of their faces. Perhaps their faces were shattered when they fell into the cave?

Naledi, like Floresiensis and Luzonensis, shares some more modern traits with other members of the homo genus, and some traits with the older australopithecines. Unlike them, we have yet to uncover evidence that Naledi used tools.

The we have a couple of unnamed hominins

These so-called “ghost populations” are known entirely from their presence in the DNA of modern humans. We don’t have any fossils from them, either because they lived in areas where the weather didn’t favor preservation, or the modern political climate makes searching for fossils difficult.

The pygmies and Bushmen derive about 2% of their DNA from an archaic population or two that we estimate split off from the rest of us about 700,000 years ago. They met and mated with these other hominins around 35,000 years ago.

More interesting is another ghost population that shows up in the genomes of west African groups like the Mende. Now, the average non-African has about 1-4% Neanderthal DNA, and Melanesians have about 4-6% Denisovan, but some tribes in west Africa, such as the Yoruba, Mende, Gambians, and Esan, may derive about 10% of their DNA from an otherwise unknown ghost population that split off before the Neanderthals! (Razib’s very nice article summarizing the paper.)

 

Oh, jeeze, it’s three am, let me finish this in the next post…