Tag Archives: Homo sapiens

African geneses: Bushmen

271-257 thousand years ago

You’ve probably run into some version of the factoid that there is more genetic variation in Sub-Saharan Africa than in all the rest of the world. This assertion has to be handled with care. It doesn’t necessarily apply to genes that have been under strong diverging selection pressures on different continents. Consider skin pigmentation: there is not more variation inside Africa than outside it in skin color, or in genes for skin color. Obviously. Likewise for hair form. But it’s true for neutral genetic variation, which is most genetic variation.

The simplest way to account for the broad Africa/non-Africa distinction would be to assume a large homogenous founder population in Africa, with a smallish number of people leaving Africa and going through a genetic bottleneck, thereby reducing their genetic variation. But recently we’ve been learning that the African situation is more complicated. Specifically, there used to be a lot of genetic differentiation between different regions within Africa. Recent population movements have smoothed out some of that variation, but recent work on ancient DNA has been bringing this more variegated past to light.

A case in point: the latest data imply that the Bushmen of Southern Africa separated from other African populations (East African, West African) around 260,000 years ago (at least), long before the major Out Of Africa venture by modern humans. What’s more, the very latest data imply that the Bushmen have received outside genetic input pretty recently, in the last 1-2 thousand years. This admixture, 9-22% of the ancestry of modern Bushmen, is absent from a 2,000 year old skeleton form Ballito Bay, South Africa.

The intruding population were probably pastoralists whose livestock, and a fraction of their genes, ultimately derived from the Near East. Another fraction of their genes originated in the Sahel or East Africa. And they probably spoke a language in the Nilo-Saharan or Afro-Asiatic family. These language families pop up as a substrate in East Africa, although largely overlain by the later expansion of Bantu speakers.

One implication: Bushman groups like the !Kung have often been presented as models for our Pleistocene hunting and gathering ancestors. Yet the most recent findings imply that there has been substantial interaction, including gene flow, between Bushmen and non-hunter-gatherers for some time.

A relevant result from twentieth century anthropology: when Nancy Howell did her classic work on the demography of the Dobe !Kung Bushmen, she found that, when you look at female reproductive histories, the Dobe !Kung look like a growing population, but when you look at male reproductive histories, they look like a shrinking population. There’s no contradiction here: the women, but not the men, in the population were sometimes having children by outsiders, neighboring pastoralists. The pastoralists in question were Bantu, having arrived in the last few centuries, but the latest genetic data imply that something similar was going long before the Bantu showed up. Since it’s not clear what effect this subaltern sexual status might have had on Bushman social organization, the social life of historic Bushmen may not be a good model for hunter-gatherer life before agriculture.

Here’s the article on Ballito Bay Boy.

And a recent review article from Nature

And assorted blog posts from Razib Khan, on the ancient Bushman split, and more recent African prehistory.


African geneses

303 – 287 thousand years ago

Our picture of human evolution in Africa around 300 thousand years ago has changed dramatically in just the last few years.

brokenhillHere’s something we already knew. This skull was found at Broken Hill, Zambia, in 1921, He (yes, “he,” he’s probably male) is sometimes known as Rhodesian Man. He looks like he’s a step away from Homo erectus, but not quite Homo sapiens. He’s heavily built, with massive brow ridges. (He looks like he could pass the “pencil test” for erectus: you could rest a pencil on those ridges. Of course, seriously, this isn’t enough to define a species.)  But he’s got a flat face and relatively large brain. He could be significantly younger than 300,000 years ago.

jebel irhoudBut now Rhodesian Man is bracketed both geographically and evolutionarily by some new finds. From Jebel Irhoud, Morocco, around 315 thousand years ago, come these skulls, which are more unequivocally Homo sapiens. pushing the fossil record of our species back 100 thousand years. The skull is still archaic ­– elongated rather than globular like a modern human – but the face is now tucked under the skull, as it is with us. The brow ridges are not as pronounced as with Rhodesian man, although still heavy for modern Homo sapiens.

naledi.jpgAnd we now have dates for Homo naledi, from South Africa, of 335-236 thousand years ago. This recently discovered species had a tiny brain, and may have been adapted for climbing trees, but still makes it into genus Homo based on other features (teeth and jaws, lower skeleton). The initial guess from a lot of folks was that this was a very early member of our genus, somewhere around early Homo erectus or earlier. But instead, Homo naledi looks to have been around at the same time as early Homo sapiens.

In other words, Africa 300,000 years ago was home to an impressive variety of humans – archaic Homo sapiens in Morocco, near relations in Zambia, and barely-humans in South Africa.

Heidelberg and Bodo

By 600 thousand years ago we’re finding people who don’t fit comfortably into Homo erectus. A jawbone from around this time was unearthed in Germany, near Heidelberg, in 1907. Another find from the same period, often assigned to the same species, comes from Bodo, Ethiopia (below).


This guy clearly isn’t modern Homo sapiens, but his brain is starting to get out of the Homo erectus range (1200 cubic centimeters cranial capacity), and his browridge is a double arch, rather than an erectus-style straight bar. He’s also got cut marks on skull and face, from someone “defleshing” him.

The exact relationship of these guys to later humans has been unclear. A popular theory assigned both to an intermediate species, Homo heidelbergensis. However yesterday we reviewed very recent genetic modeling by Rogers and coworkers that suggests that a three-way split between Homo sapiens, Neanderthals, and Denisovans has already happened by this point, although the lineages haven’t had long to differentiate. So probably Heidelbergers in Europe are ancestral to Neanderthals (or at least closely related to their ancestors), while Bodo man is similarly related to Homo sapiens.

The Inheritors

42.4-40.2 thousand years ago

“The Inheritors” is a novel by William Golding about the encounter between Neanderthals and modern humans. Like another Golding novel, “Lord of the Flies,” it is written with a mid-twentieth century awareness that advanced societies don’t leave behind the potential for cruelty. The novel isn’t all that scientifically accurate, though: Golding’s early humans have bows and arrows, for example.

Neanderthal 1, the first Neanderthal fossil recognized as probably belonging to another species, was discovered in the Neander Thal (=Neander Valley) in 1856. He is close in time to the last Neanderthals: the most recent review of the evidence finds that Neanderthals disappear as a distinct group around 40 thousand years ago. They were almost certainly outcompeted by Homo sapiens who had arrived in Europe earlier. There was probably some kind of coexistence between Neanderthals and H. sapiens over many thousands of years. Regional cultures from this period, like the Châtelperronian, may represent Neanderthals copying the Aurignacian culture of incoming H. sapiens. The final blow may have come between 39 and 38 thousand years ago, when a “Heinrich event” sent cascades of icebergs into the North Atlantic, drastically chilling Europe. Homo sapiens recovered from the cold spell; Neanderthals did not.

Just last year we have learned that one of the earliest modern human fossils from Europe, Oase 1, from Romania (40 kya), has substantially more Neanderthal DNA, 6-9%, than living Europeans. Furthermore, this DNA comes in the form of long stretches of chromosomes, rather than little bits broken up by millennia of recombination, showing that his Neanderthal ancestors go back just a few generations, maybe to some great-great-great-grandparents. We’ve also learned from isotopic evidence that Oase 1 got the proteins in his diet from a broad array of sources, including freshwater fish.

And here is some of the art produced by Homo sapiens 40 thousand years ago. lionman The lion man from Hohlenstein-Fels

And the earliest known cave paintings, from Sulawesi, Indonesia. indonesiancaveart


We have been treating Neanderthals here as a species, Homo neanderthalensis, distinct from our own species, Homo sapiens. Some researchers elect to call Neanderthals a subspecies, Homo sapiens neanderthalensis, and classify modern humans as another subspecies, Homo sapiens sapiens.

The line between subspecies and species is not clear cut, nor – given the way evolution works – should we expect it to be. Recent work on ancient DNA recovered from fossils has shown just how complicated the subject is. The spectacular finding of the last few years is that modern humans are hybrids, getting most of their ancestry from a single founding population (we can call them Homo sapiens), but incorporating limited ancestry from close relatives. Thus human beings outside Africa have 1-4% Neanderthal DNA. So it looks as if early in the course of expansion(s) out of Africa, there was a limited amount of interbreeding with Neanderthals.* And not just with Neanderthals. Populations in Melanesia get an additional 4-6% of their DNA from a widespread East/Southeast Asian population known as Denisovans, while some African groups may have ancestry from non-sapiens populations in Africa. (The fossil record for Denisovans is a lot sparser than for Neanderthals, and it’s even sparser for African non-sapiens.)

This isn’t reason enough to put Neanderthals and sapiens in a single species: plenty of species occasionally hybridize with related species. And in fact the DNA evidence implies that sapiens and Neanderthals were moving toward being reproductively isolated. Specifically, we find that a lot of Neanderthal genes related to testis development and male fertility are underrepresented (i.e. at a lot less than 1-4% frequency) in modern humans. The likely explanation is that those genes didn’t work well against a H. sapiens genetic background. In other words, if you were mixed sapiens/Neanderthal man, you probably had fertility problems, albeit not to the point of complete sterility. On the other hand, other Neanderthal genes  – especially genes related to immune function – were useful to modern humans moving into Neanderthal territory and are found at high frequency in Europeans today.

There is an extensive older literature in physical anthropology on “race crossing.” Researchers were concerned with whether people with mixed racial ancestry might have reduced fitness as a result of combining incompatible genes. This literature is reviewed at book length here. The overwhelming evidence is that “race crossing” has no harmful biological consequences (in contrast to close inbreeding, which is a bad idea: check out this post on the Habsburgs.)

Some new data put this in perspective. My colleagues Alan Rogers, Chad Huff and Ryan Bohlender have just shown that the population ancestral to Neanderthals and  Denisovans separated from the ancestors of Homo sapiens somewhat more than 750,000 years ago, passing through a narrow population bottleneck, probably in an early Out Of Africa episode (before the later modern human Out Of Africa episode). This population then split somewhat less than 750,000 years ago into Neanderthal and Denisovan branches. So Homo sapiens and Homo neanderthalensis evolved separately for the better part of a million years, and were some way on the path to reproductive isolation. By contrast, different populations (“races”) within Homo sapiens have been evolving separately for 100,000 years or less outside Africa, and perhaps 250,000 years within Africa. This has been enough time to evolve major differences in traits like skin color and hair form, but apparently not to create appreciable biological barriers to interbreeding.

And here’s a link covering some recent research suggesting that across a wide range of organisms it takes a surprisingly clock-like average of two million years separation to split one species into two.

* Hence, my Neanderthal name is Carg, and my website is cargshome.

Australian megafauna and the Sixth Great Extinction

The Earth has been through a number of mass extinctions. On Logarithmic History, we covered the five greatest mass extinctions during the month of March, and into early April.

For all these extinctions, the most likely cause is some kind physical catastrophe: either drastic changes in the chemistry of oceans and atmosphere, or extraterrestrial events, like asteroid strikes or, just possibly, gamma ray bursts.

Competition with other organisms is another major cause of extinctions. Usually this is just part of a steady background level of extinction. But occasionally competition – and extinction rates – increase dramatically, as during the Great American Interchange five million years ago, when South American animals were swamped by North American invaders with the establishment of the Isthmus of Panama.

Earth now may be on the edge of another episode of mass extinction, a Sixth Extinction. (although rates of species extinction don’t match those of earlier ME’s). This time the cause is very different: a single species, Homo sapiens, is playing an overwhelming role. Although the pace of extinction has accelerated over the last few centuries, you can make a case that the sixth extinction began a long time ago, with the expansion of modern humans out of Africa. Australia in particular sees the disappearance of a unique fauna that evolved over more than a hundred millions of years of isolation. This fauna included monster wombats, giant kangaroos, huge flightless birds, and a marsupial version of a lion.

All of these – all land mammals, reptiles, and birds with mass of more than 200 pounds – seem to have gone extinct by around 45,000 years ago, after humans crossed the sea to settle the island continent. (Sea levels were lower then, and Australia was connected with New Guinea, but still isolated from mainland Asia.) This is not settled science. Some researchers think climate change was to blame. But I think the evidence in the case of this and later mass extinctions is strongly in favor of humans as the major agent of extinction. This is one more reason to treat the advent of our species as one of the major evolutionary transitions, comparable to the evolution of complex cells, or multi-cellular life.

Two roads diverged

53-50 thousand years ago.

The broad outlines of the spread of Homo sapiens have been established for several decades now: origins in Africa, expansion out of Africa at least 50-45 thousand years ago. But we’re still arguing about the details. Just this year it’s been reported that modern humans reached Sumatra 73-63 thousand years ago, and that stone tools in Australia date back 60 thousand years ago.

Also, based on recent recalibrations of DNA mutation rates, it looks like the African/non-African split might have happened more like 100 thousand years ago than 50 thousand years ago. So the ancestors of non-African (or non-sub-Saharan-African) H. sapiens might have occupied a homeland somewhere north of the Sahara between 100 and 50 thousand years ago, before spreading through Eurasia. North Africa is one possibility. The Near East, maybe the Arabian peninsula, is another possibility. The (or “a”) homeland might be (gated, sorry) underwater, under the Persian Gulf (sea levels were lower then). Both possibilities have some archeological support. There might have been multiple homelands, and multiple expansions – south through Arabia and along the shores of the Indian Ocean, and north through the Levant and into Europe.

A recent (2015) redating of archeological finds suggests that the Levant-to-Europe corridor was part of the story. A modern stone tool technology, coming from Ksar Akil, just outside Beirut, Lebanon, dates to about 50,000 years ago, a little before much the same technology appears in Europe, in the form of the Upper Paleolithic.

And many other “details” remain to be resolved: What did interbreeding with non-sapiens mean for the evolution of H. sapiens? And just what advantage(s) did H. sapiens have that allowed him (us!) to replace other species? Stay tuned for more on Logarithmic History