The end of Eden
A particular conception of the “Out of Africa” model of human origins died in this decade. This model hooked into preexistence narratives about “Adam” and “Eve”, utilizing Y and mitochondrial DNA lineages passed down through direct male and female lines respectively. Its most extreme manifestation could be exemplified by Richard Klein’s ideas in the early 2000s outlined in his book .
For Klein the chasm between Homo sapiens sapiens, humans, and other hominins was vast. A physical anthropologist who surveyed with skill the rapid expansion and proliferation of modern human cultures over the past ~50,000 years, Klein relied on a particular evolutionary model to explain how this occurred. He posited that humankind emerged in East Africa as a punctuated speciation event, triggered by a mutation which allowed for the development of fully elaborated recursive language.
The difference between our own lineage and our relatives in this framework was huge. To not put too fine a point on it, Neanderthals and other archaic humans were animals. We, Homo sapiens sapiens, were humans qua humans.
Though Klein was a paleoanthropologist, he gained great support from a school of molecular evolution which arose in the 1970s and 1980s under Allan Wilson. Wilson’s initial fame arose because he utilized a “molecular clock” analysis of primates to contend that the divergence of our human lineage from great apes was much more recent than paleontologists had believed. Eventually new fossil finds confirmed the molecular phylogeny. After this event Richard Leakey has stated paleoanthropologists were reluctant to challenge molecular results.
Wilson later focused on recent on human origins, utilizing mitochondrial DNA, which is passed down directly through the maternal lineage. In this way they found that African mtDNA lineages were very diverse, and that non-African lineages were nested within the broader tree of African lineages.
The conclusion from this finding was that modern humans arose in Africa and spread to other parts of the world. This conclusion in general has been confirmed.
But over the years more and more evidence has accumulated that the story is more complicated than the original narrative that all modern humans descend from a small bad of East Africans who populated the whole world ~50,000 years ago.
Dissenters from Eden
There were always geneticists who were skeptical of the neat Out of Africa with total replacement model. In Richard Leakey recounts a conference in 1992 where he was pigeon-holed by geneticists who thought there was no reason to accept without dispute the mitochondrial Eve narrative. Over the years talking to some older geneticists I can say that Leakey was reporting a real undercurrent of irritation with the confidence that Allan Wilson’s group and their fellow travelers projected in relation to their model. Nordborg 1998 On the probability of Neanderthal ancestry reflects some of the technical objections to inferring too much from one locus when it came to the possibility of other components of ancestry.
When genome-wide analyses in the middle 2000s became feasible, a visible counter-culture within genetics argued that total replacement was not supported by the data. In 2006 Wall & Hammer published Archaic admixture in the human genome. They concluded that “Recent work suggests that Neanderthals and an as yet unidentified archaic African population contributed to at least 5% of the modern European and West African gene pools, respectively.” They were not that far off with European populations. As far as Africa goes, that is a question that will be explored in detail in the next few years.
That analysis though has only 44 citations. I have had debates on Twitter with how exotic and marginal these ideas were. In general it is safe to say that they were not exotic and marginal in the community of human evolutionary population geneticists. But that’s not a large set. Both and Milford Wolpoff have indicated a lot of marginalization for models outside of the narrow window of Out of Africa with total replacement. From everything I’ve heard about the run up to the 2010 publication of the Neanderthal genome many of the principal researchers, including Svante Paabo, were totally surprised by the evidence of admixture into modern lineages. Wolopff even emailed me after I reviewed the paper to suggest that it felt so good to come out of the wilderness and have some of his views accepted.
Anagenesis and punctuation?
But were Wolpoff’s views accepted? The revised model actually kept much of the Out of Africa framework in place, except it added the wrinkle of assimilation of some archaic lineages. The dominant signal in the non-African genomes seems to have come from an African lineage which left around ~50,000 years ago.
The classical multi-regional model that Wolpoff was associated with, whereby modern humans evolved across the whole world from local archaic lineages, but maintained species cohesion through gene flow, was not supported. Rather, the archaic admixture of Neanderthals and Denisovans into Oceanians pointed to local continuities, which was a broader position of multi-regionalism. But this is not speciation without branching, anagenesis.
Nevertheless, there was another aspect of Out of Africa with replacement that needed revision. Though not explicitly outlined in many framings, one aspect implicit is that the dynamics that Africa and Eurasia were subject to during the emergence of modern humans were the same.
But that doesn’t seem to be the case. The ancestors of all non-Africans went through a major population bottleneck. On the order of ~1,000 individuals (this is a very large bottleneck actually, and I’ve seen numbers as low as 100, though that seems on the small side; calculating effective population size ~50,000 years ago can be tricky). The same is not true of African populations. Though many of them show signals of population declines during the Pleistocene, the extreme uniform bottleneck which characterizes all non-Africans, from Iberia to Australia to Patagonia is just not evident in Sub-Saharan African populations.
In other words, the Out of Africa event did not apply within Africa. Here’s an excerpt of an email I sent to Carl Zimmer in December of 2010 (he was updating the second edition of ):
…it may be that there was no rapid antique population expansion in Africa which was analogous to [the] out of Africa migration. IOW, non-Africans are just a branch of Northeast Africans, and the Bushmen and other groups were already differentiated by that point. So you could theoretically remove the arrows within Africa! I think this is a subtle and tendentious point, so probably best to leave that as it is. But remember how deep the basal branching of the Bushmen was in the Denisova paper? It WAY predates any possible out of Africa migration by multiples.
Which brings me to the current year and the present time. The recent paper which utilized an ancient genome from South Africa to push back the date of the diversification of African lineages to about ~250,000 years before the present was not entirely surprising to me. Every time I talked to people who had access to African whole genomes their dates kept getting pushed back further and further into the past.
And of course we now have fossil confirmation that human populations which seemed to be anatomically modern (or close) were already present ~300,000 years ago in Morocco. The New York Times has a good overview of the work, Oldest Fossils of Homo Sapiens Found in Morocco, Altering History of Our Species. I read the papers and the commentaries and don’t have much to add, nor do they add much for non-specialists in my opinion (since we can’t really judge the morphology too well, nor do we have a detailed understanding of the fossil record). In one of the Nature letters the authors conclude in the abstract that “The emergence of our species and of the Middle Stone Age appear to be close in time, and these data suggest a larger scale, potentially pan-African, origin for both.”
This suggest to me anagenesis. Has multi-regionalism come back, but no within Africa?
Parameters, not paradigms
John Hawks has put in his two cents, and it’s always worth paying attention. My major take home is that we don’t know a lot even though we know more, and we need to be careful here. The genome blogger from the 2000s who has been relatively quiet over the last five years, Dienekes, resurfaced, dismissing the idea of pan-African anagenesis and asserting an Out of North Africa viewpoint. He’s been talking about this model since 2011, so there’s nothing new here. In January of 2011 he asserted that “Africa was home to a structured population.” That is what we are seeing today.
The publication of the Nature letters triggered a lot of discussion on Twitter. When I was involved it mostly consisted of Aylwyn Scally and Pontus Skoglund, with John Hawks, Chris Stringer, and others jumping into the stream. Here are some points which are of note:
1) Most people now suspect that large scale population structure within Africa over the past few hundred thousand years is a major story.
2) But there is an assumption that collapsing of that structure through gene flow was not reciprocal. That is, some populations likely expanded at the expense of others. The arguments are whether the assimilation of the secondary groups is on the order of a few percent, as seems to be the case in Eurasia, or a much higher fraction.
3) Because the phylogenetic distance between within African lineages is likely smaller than between Neanderthals and modern humans, as well as the likely similar census sizes and technological toolkits, I contended that it is not unreasonable to guess that as much as 20% of the ancestry of a daughter population of an expanding group could be from the local substrate. There was no great objection to this guess.
4) Remember, even simple mtDNA phylogenies as far back as the 1980s, as well as paleontological analyses of fossils, indicated that an Out of Africa movement into Eurasia. This was such a strong signal in the data that it was clear with even relatively little to go on. The situation for within Africa is not analogous, suggesting to me that an extreme model of replacement or gene flow across persistent demes in local regions is not tenable.
5) Ultimately, the issue will resolve on parameters of admixture and the nature of demographic expansion in the details. Instead of a tree, we will conceive of this as a graph, a trellis with lengths of different thicknesses.
6) Hawks brought up the fact that one reason classic multi-regionalism did not work is that the Fisher wave of expansion of favored genes is slower than the migration of humans. When I suggested it does not seem that the genetics of gene flow in plants, which do resemble classical multi-regionalism, were a good analogy for humans, Skoglund contrasted the sessile nature of the taxon in contrast to mobile humans. I did point out though that after favored alleles moved through migration into a population, there was often in situ selection. He agreed.
7) A key issue that both Hawks and Dienekes emphasize is that we don’t know the role that extremely diverged lineages from our own ancestors play in our story. That is, were there many modern human populations across Africa, interspersed with other human species? Or was there one modern human population that mixed with other species? We don’t really know the details of all of this.
8) I expressed skepticism of the idea of “behavioral modernity.” My reason for being skeptical is that the origin of modern humans is not as neat as we like to think, and the origin of “behavioral modernity” is also not as neat as we like to think. When the consensus was that humans emerged as a punctuated de novo event, ensouled by the Lord God on High 50,000 years ago (or, coming down from the skies as in or in Larry Niven’s ), the idea of behavioral modernity kind of made sense. But it’s all more confused now (in any case, in Clive Finlayson’s he seems to be arguing that much of modern culture was invented by Gravettians, well after the Out of Africa event).
The consensus seems to be that rather than focusing on a set of human universals as behaviorally modern, we should look at the demographic patterns of the past to infer when our own lineage came into its distinctive being. Those of you who have read me for a while know this is already congenial to me, Luke Jostins’ plot of the encephalization of all hominin lineages over the past million years was suggestive to me long ago that our own lineage is not so special. Rather, something like us was probably inevitable so long as an asteroid didn’t wipe out large mammals once Homo erectus spread across the globe. Humanity is a destiny, not a lineage.