Dienekes has a post up highlighting a preprint out of Pontus Skoglund’s group. It is titled Ancient genomes mirror mode of subsistence rather than geography in prehistoric Europe. It doesn’t seem to be online (fingers crossed that it shows up linked at Haldane’s Sieve soon). In any case I am not surprised by the broad outlines of the thesis. And, it is not as if Skoglund’s group is the only one working in this area, I have suspicions that others are finding something very similar. These results out of Europe are probably reflective of the fact that much of the model in Peter Bellwood’s First Farmers is generally correct, the emergence of an agriculture revolution in a few select world societies produced a cultural and demographic revolution.
Over at Scientific American Eric Michael Johnson has a very long post up, The Case of the Missing Polygamists. It is a re-post of something he already published at Psychology Today a few years ago. Though provisionally a review of Sex at Dawn, Johnson covers a lot of ground, and also has extensive quotations from Sarah Blaffer Hrdy.
I’m reflecting upon the post for a second time because it is very rich in ideas, and lays out may different general concepts and specific propositions. The bottom line from what I can gather is that Johnson agrees with those thinkers who believe that agriculture and the Neolithic revolution to a great extent reshaped social relations, and give us a skewed perception of “normal” human societies. I’m not going to rehash all of the points in the piece, but will focus on just a few which I think I can extend upon fruitfully.
The figure to the left is from a new paper in Science, When the World’s Population Took Off: The Springboard of the Neolithic Demographic Transition. It reports the findings from 133 cemeteries in the northern hemisphere in regards to the proportion of 5-19 year old individuals. When calibrated to period when agriculture was introduced into a specific region there seems to be a clear alignment in terms of a demographic transition toward a “youth bulge.” Why? A standard model of land surplus explains part of it surely. When farmers settle “virgin land” there is often a rapid “catch up” phase toward the Malthusian limit, the carrying capacity. Another possibility though is that sedentary populations did not need to space their offspring nearly as much as mobile hunter-gatherers. Whatever the details, the facts remain that the data do point to a shift in the age pyramid during this period. The author wonders as to the possible cultural implications of this. There is an a priori assumption that a young vs. old age profile in a society constrains its choices and channels its energies (e.g., think the “baby boom” generation in the USA). A final interesting point is that the authors note that today we are seeing the last gasp of this transition toward large numbers of children, as fertility drops toward replacement all across the world. That too may have some cultural consequences.
Seriously, sometimes history matches fiction a lot more than we’d have expected, or wished. In the early 2000s the Oxford geneticist Bryan Sykes observed a pattern of discordance between the spatial distribution of male mediated ancestry on the nonrecombinant Y chromosome (NRY) and female mediated ancestry in the mitochondrial DNA (mtDNA). To explains this he offered a somewhat sensationalist narrative to the press about possible repeated instances of male genocide against lineage groups who lost in conflicts.
Here is a portion of the book of Numbers in the Bible:
15 – And Moses said unto them, Have ye saved all the women alive?
16 – Behold, these caused the children of Israel, through the counsel of Balaam, to commit trespass against the LORD in the matter of Peor, and there was a plague among the congregation of the LORD.
17 – Now therefore kill every male among the little ones, and kill every woman that hath known man by lying with him.
18 – But all the women children, that have not known a man by lying with him, keep alive for yourselves.
Then there is the rape of the Sabine women. The ethnogenesis of the mestizo and mulatto populations of the New World in large part was the union between non-European women and European men. These are hard brutal myths and hard brutal facts. But do they reflect an essential aspect of the dynamics which have shaped our species’ past?
I’m not willing quite yet to add a confident weight upon this possibility, but this seems to be part at least part of the picture. You see a major disjunction on male and female lineages among South Asians for example. A new paper in PNAS adds weight to this possibility, albeit only incrementally. Ancient DNA reveals male diffusion through the Neolithic Mediterranean route:
The Pith: Over the past 10,000 years a small coterie of farming populations expanded rapidly and replaced hunter-gatherer groups which were once dominant across the landscape. So, the vast majority of the ancestry of modern Europeans can be traced back to farming cultures of the eastern Mediterranean which swept over the west of Eurasia between 10 and 5 thousand years before the before.
Dienekes Pontikos points me to a new paper in PNAS which uses a coalescent model of 400+ mitochondrial DNA lineages to infer the pattern of expansions of populations over the past ~40,000 years. Remember that mtDNA is passed just through the maternal lineage. That means it is not subject to the confounding dynamic of recombination, allowing for easier modeling as a phylogenetic tree. Unlike the autosomal genome there’s no reticulation. Additionally, mtDNA tends to be highly mutable, and many regions have been presumed to be selectively neutral. So they are the perfect molecular clock. There straightforward drawback is that the history of one’s foremothers may not be a good representative of the history of one’s total lineage. Additionally the haploid nature of mtDNA means that genetic drift is far more powerful in buffeting gene frequencies and introduced stochastic fluctuations, which eventually obscure past mutational signals through myriad mutations. Finally, there are serious concerns as to the neutrality of mtDNA…though the authors claim to address that in the methods. I should also add that it also happens to be the case that there is less controversy and more surety as to the calibration of mutational rates of mtDNA than the Y chromosomal lineages of males. Their good for determining temporal patterns of demographic change, and not just tree structures.
Here’s the abstract, Rapid, global demographic expansions after the origins of agriculture:
About five months ago I read Peter Bellwood’s First Farmers: The Origins of Agricultural Societies. Bellwood’s thesis is simple: that the first adopters of farming entered into a period of rapid demographic expansion and by and large replaced non-farming groups. The populations which dominate the world today in this model are then the descendants of the very small set of cultures which ~10,000 years ago triggered the Neolithic Revolution. When Bellwood presented his thesis in the mid-2000s many would have dismissed it out of hand. Today I believe we have to take this model seriously.
There are two primary reasons from my perspective why I am now thinking about Bellwood’s thesis a great deal. First, the archaeogenetic inferences based on distributions of modern allele frequencies which suggested that the Neolithic Revolution in Europe was a matter of cultural diffusion seem far shakier. With such genetic models no longer taken for granted the recent historical, semi-historical, and ethnographic evidence, on farming transitions must be given much more weight. The case of the Bantu expansion in Africa seems to be semi-historical. The Bantu farmers themselves were not literate but their wave of advance was in historical time. Tellingly, the Bantu speaking populations of Southern Africa are genetically more similar to the Fang of Cameroon then they are to the Khoisan to their west! More well documented has been the attempts by Europeans to settle various lands overseas in their colonial adventures. They have been able to marginalize populations which did not habitually practice intensive agriculture relatively easily (note that the locus of Afrikaner settlement was initially around the Cape, where Bantu influence was minimal and Khoikhoi pastoralists were dominant). In contrast, in regions like Mesoamerica where obligate intensive agricultural civilization had deep roots there was no biological replacement, but hybridization.
An argument can be made that the initial farmers did not have so many advantages over their hunter-gatherer neighbors. So the power and force of the mass agricultural way of life which bore down upon the indigenes of Australia was qualitatively different because the Europeans who arrived were the outer wave of an ancient and ruthlessly efficient civilization of farmers, honed to brutal perfection through the cauldron of inter-group competition over thousands of years. I think the best counter argument against this is the evidence of rapid sweeps of cultural forms in European in prehistory, as well as rapidity of the success of the Bantu agricultural toolkit.
The new genome blogger Diogenes expresses the thesis of agricultural replacement to near maximal levels in the model which he is attempting to test with ADMIXTURE runs. Here are his propositions (formatting reedited for clarity):
The economist Samuel Bowles recently had a paper out in PNAS which caught my attention, Cultivation of cereals by the first farmers was not more productive than foraging. This naturally begs the question: why did farming conquer foraging as a lifestyle? First, let’s look at the abstract:
Did foragers become farmers because cultivation of crops was simply a better way to make a living? If so, what is arguably the greatest ever revolution in human livelihoods is readily explained. To answer the question, I estimate the caloric returns per hour of labor devoted to foraging wild species and cultivating the cereals exploited by the first farmers, using data on foragers and land-abundant hand-tool farmers in the ethnographic and historical record, as well as archaeological evidence. A convincing answer must account not only for the work of foraging and cultivation but also for storage, processing, and other indirect labor, and for the costs associated with the delayed nature of agricultural production and the greater exposure to risk of those whose livelihoods depended on a few cultivars rather than a larger number of wild species. Notwithstanding the considerable uncertainty to which these estimates inevitably are subject, the evidence is inconsistent with the hypothesis that the productivity of the first farmers exceeded that of early Holocene foragers. Social and demographic aspects of farming, rather than its productivity, may have been essential to its emergence and spread. Prominent among these aspects may have been the contribution of farming to population growth and to military prowess, both promoting the spread of farming as a livelihood.
My own working assumption is that the “first farmers” existed in a state of land surplus, and so like the medieval peasants in the wake of the Black Death found themselves released from Malthusian constraints, at least until their natural increase swallowed up their affluence. Bowles gives several reasons to be skeptical of this conjecture. The list in table 2 shows the positive and negative biases in the model when one back-projects later stages of farming to the initial period. Metal tools, well developed distribution channels, and more productive varieties, were features of mature agricultural societies. On the time-insensitive scale the necessity of planning ahead and waiting patiently for the crop to be ripe count against the gains of the farming way of life as well. The main variable which would weight in favor of farming is the land surplus alone, though Bowels argues that the ethnographic data as to the benefits of a surfeit of this input factor of production is mixed. I am skeptical of this point, though I can’t say I’ve dug deeply into the literature.
There are other factors, such as the fact that farmers were immobile, and so subject to attack. A shift toward a few crops also reduced the diversity of the diet, and therefore entailed a trade off between a diet rich in micronutrients, fiber, protein, and fat, to one overloaded on carbohydrates. Finally, a reliance on a few crops also means heightened “tail risk.” Think of the Irish potato blight. Hunter-gatherer populations would usually have a more diverse portfolio, and so be buffered more from environmental shocks.
After linking to Marnie Dunsmore’s blog on the Neolithic expansion, and reading Peter Bellwood’s First Farmers, I’ve been thinking a bit on how we might integrate some models of the rise and spread of agriculture with the new genomic findings. Bellwood’s thesis basically seems to be that the contemporary world pattern of expansive macro-language families (e.g., Indo-European, Sino-Tibetan, Afro-Asiatic, etc.) are shadows of the rapid demographic expansions in prehistory of farmers. In particular, hoe-farmers rapidly pushing into virgin lands. First Farmers was published in 2005, and so it had access mostly to mtDNA and Y chromosomal studies. Today we have a richer data set, from hundreds of thousands of markers per person, to mtDNA and Y chromosomal results from ancient DNA. I would argue that the new findings tend to reinforce the plausibility of Bellwood’s thesis somewhat.
The primary datum I want to enter into the record in this post, which was news to me, is this: the island of Cyprus seems to have been first settled (at least in anything but trivial numbers) by Neolithic populations from mainland Southwest Asia.* In fact, the first farmers in Cyprus perfectly replicated the physical culture of the nearby mainland in toto. This implies that the genetic heritage of modern Cypriots is probably attributable in the whole to expansions of farmers from Southwest Asia. With this in mind let’s look at Dienekes’ Dodecad results at K = 10 for Eurasian populations (I’ve reedited a bit):
When it comes to the synthesis of genetics and history we live an age of no definitive answers. L. L. Cavalli-Sforza’s Great Human Diasporas would come in for a major rewrite at this point. One of the areas which has been roiled the most within the past ten years has been the origin and propagation of the agricultural lifestyle across the European continent between 10,000-6,000 years before the present (starting in Europe’s southeast fringe a few thousand years after the origination of the Neolithic lifestyle in the Levant, and finally pushing into the southern Scandinavian peninsula only ~6,000 years ago). The reasons for this particular debate about the origin of the European are manifold. First, most scholars are of European ancestry, and some of the debates have roots going back a century. So a natural interest exists based on normal human biases. Second, when it comes to genetics the climate of Europe is ideal for the preservation and extraction of ancient DNA. Third, there are relatively clear and distinct theoretical models which can be tested by the data, whether to verify or refute.
I have already reviewed earlier work in three previous posts, European man perhaps a Middle Eastern farmer, European man perhaps not a Middle Eastern farmer, and Völkerwanderung back with a vengeance. Instead of rehashing everything I’ll take it as a given that you’ve read or skimmed those posts. Rather, let’s move on to a new paper in PLoS Biology, Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities:
The cockroach as we know it has been around for ~140 million years. That’s a rather long run. The evolutionary design of the cockroach seems to be well suited to avoiding obsolescence; it’s withstood the test of time. I suspect that the particular example of the roach is often used to illustrate the blindness of evolution because of its lack of aesthetic alignment with the the values of modern humanity. Unlike the elegant wasp or the industrious bee the cockroach seems to have few redeeming characteristics on first blush. The Hutus referred to the Tutsis as cockroaches before and during the Rwanda genocide of 1994. And yet the roach succeeds, it breeds, and it flourishes.
Some of the same class of issues pertain to our own species. What we feel to be edifying, to be aesthetically pleasing, may not comport with the final judgement of history, of evolution. The narrative of man ascending which has become so popular since the Enlightenment turns out to present us with some problems when one realizes that our species seems to have regressed on particularly transparent metrics such as height and cranial capacity since the last Ice Age. But the prevailing wisdom of the ancients that we descend from an Edenic Golden Age also does not seem to necessarily comport with the record at hand either. Just as the past is cloudier than we once perceived it to be, so the future often looks muddled from the perspective of the present. How did man come to be? What should we be? And why should we be? These are a combination of positive and normative questions, and Spencer Wells tackles them in his newest book, Pandora’s Seed: The Unforeseen Cost of Civilization.