43,500-year old Aurignacian north of the Alps

PNAS doi: 10.1073/pnas.1412201111

Early modern human settlement of Europe north of the Alps occurred 43,500 years ago in a cold steppe-type environment

Philip R. Nigst et al.

The first settlement of Europe by modern humans is thought to have occurred between 50,000 and 40,000 calendar years ago (cal B.P.). In Europe, modern human remains of this time period are scarce and often are not associated with archaeology or originate from old excavations with no contextual information. Hence, the behavior of the first modern humans in Europe is still unknown. Aurignacian assemblages—demonstrably made by modern humans—are commonly used as proxies for the presence of fully behaviorally and anatomically modern humans. The site of Willendorf II (Austria) is well known for its Early Upper Paleolithic horizons, which are among the oldest in Europe. However, their age and attribution to the Aurignacian remain an issue of debate. Here, we show that archaeological horizon 3 (AH 3) consists of faunal remains and Early Aurignacian lithic artifacts. By using stratigraphic, paleoenvironmental, and chronological data, AH 3 is ascribed to the onset of Greenland Interstadial 11, around 43,500 cal B.P., and thus is older than any other Aurignacian assemblage. Furthermore, the AH 3 assemblage overlaps with the latest directly radiocarbon-dated Neanderthal remains, suggesting that Neanderthal and modern human presence overlapped in Europe for some millennia, possibly at rather close geographical range. Most importantly, for the first time to our knowledge, we have a high-resolution environmental context for an Early Aurignacian site in Central Europe, demonstrating an early appearance of behaviorally modern humans in a medium-cold steppe-type environment with some boreal trees along valleys around 43,500 cal B.P.

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Complex Y chromosome structure in East Tyrol (and more IE thoughts)

Cultural diversity can disappear in a few generations, but genetic diversity -barring major genocides or disasters- usually persists.

The East Tyrol region in Austria has been Germanic-speaking since the Middle Ages, but historical evidence documents the presence of Romance, Germanic, and Slavic groups in its territory. How can we untangle the origin of the different groups when they are all jumbled up together now, and all Germanic-speaking? Previous research has shown that patrilineal groups can be isolated on the basis of surnames, but in the case of East Tyrol, the wide adoption of surnames happened after the region had become linguistically Germanic.

The authors of the new paper exploited the structure of local toponyms, in particular pasture names. The figure on the left shows the concentrations of Slavic (panel A), Romance (panel B), and Germanic (panel C) pasture names. While Germanic pasture names are evenly distributed, there is a contrast between those of Slavic and Romance origin. From the paper:

From the 853 analyzed pasture names in East Tyrol 71% were derived from Germanic (Bavarian) etymons, 17% from Slavic etymons, and 12% from Romance etymons. While pasture names with Germanic etymons were evenly distributed in high density within the whole study area the names with Slavic etymons were spatially focused in the east and north of East Tyrol (Fig. 2). Geographically, these are the lower Drau, Isel, Kals, Virgen and the Defereggen valleys (Fig. 1). No names with Slavic etymons were found in the southwestern Puster valley (Fig. 2). The pasture names with Romance etymons focus mainly in the southern part of East Tyrol (Gail, Puster, and Villgraten valley, Fig. 2). The slight northeastward trend observed in the distribution of Romance etymons is solely caused by the Kals valley, a medieval Romance linguistic enclave, which was separated from the Romance main territory in the 10th century [36]. On the basis of these results, East Tyrol was divided into two regions of former Romance (Puster, Gail, and Villgraten valley; region A) and Slavic (Isel, lower Drau, Defereggen, Virgen, and Kals valley; region B) main settlement (Fig. 2).

The authors dissected the occurrence of different haplogroups in the two contrasting regions (A: Romance, and B: Slavic) in some great detail:

Splitting the East Tyrolean population sample into regions A and B resulted in a partitioning of haplogroups E-M78, R-M17, R-M412/S167*, and R-S116*. E-M78, R-M17 and R-S116* Y chromosomes were exclusively found in region B whereas samples assigned to R-M412/S167*, R-U106/S21, and R-U152/S28 reached higher frequencies in region A (Fig. 3, Table S7). When attributing the samples to the fathers' and grandfathers' places of birth/residence, as reported by the participants, practically identical patterns were obtained for most of the haplogroups (Fig. 3). 

Y chromosomes belonging to haplogroups G-P15, I-M253, and J-M304 showed much lower regionalization in their frequencies (Fig. 3) at all three generation levels.

The non-localization of the G-P15, I-M253, and J-M304 seems reasonable as these may represent what is common in these populations (and one could indeed speculate -on the basis of current ancient DNA knowledge- that they correspond to Neolithic, Paleolithic, and Bronze Age processes respectively)

Two of the most interesting findings are:

Haplogroup R-M412/S167* was found at low frequencies in the combined East Tyrolean sample. However, the R-M412/S167* chromosomes were sorted by the subdivision of the study area and reached in region A levels of ~14% whereas their frequency in region B was well below the 5% threshold. At the probands and fathers level of analysis region A featured approximately fourfold higher frequencies of these chromosomes than region B. This ratio changed to about nine when placing the samples at the grandfathers' places of birth/residence. These contrasts remained statistically significant after correcting for multiple comparisons [22] at the fathers and grandfathers analysis level.

and:

The western border of the geographic expansion of haplogroup R-M17 Y chromosomes is to be found in Central Europe and largely follows the political border separating present-day Poland (57%) and Germany (East: ~30%, South: ~14%, West: ~10%) [42]. Frequencies of about 15% and 10% were also found for Austria [18] and North-East Italy [48], respectively. In South Italy and in West Europe R-M17 chromosomes are not present at informative frequencies. 

In this study, the proportion of Y chromosomes carrying the derived M17 allele was 14.1%, a value that nearly perfectly matched those reported for West Austria (North Tyrol, 15.4%) and South Germany (Munich; 14.3%) [18], [42]. However, haplogroup R-M17 was completely absent in the East Tyrolean sub-sample from region A, but made up to 16% in region B. This result remained practically unchanged when assigning the probands to their respective fathers' or grandfathers' places of birth/residence (Fig. 3).

The new study reinforces my belief that R-M17 was not originally present in the Italo-Celtic branch of Indo-European. Together with the paucity of the same lineage in Albanians (~5%), Armenians (less than 5%), and its quite uneven distribution in Greeks, it is becoming increasingly clear that R-M17 may represent a late entrant that affected minimally southern and western Europe.

The fountain of its spread was probably a trans-Caspian (?) Central Asian staging point that followed a counter-clockwise route into Europe that spawned the northern (Germanic and Balto-Slavic) groups of Europe and the Indo-Iranians, who remained longer in their BMAC homeland, finally breaking down during the 2nd millennium BC. This would also harmonize with the increasing evidence for complementary R-M17 distributions in Europe and Asia, associated with the Z93 marker. 


It might appear that Z93+ chromosomes may track the later expansion of the Indo-Iranian world. I have observed before that R-M17 seems distributed in a long arc north and east of the Caspian, and it is perhaps in different points along this arc that the dominant European (NW) and Asian (SE) types emerged out of the early Neolithic population.

Combining this insight with an analysis of Y chromosome variation within the Graeco-Armeno-Aryan group, it appears that Graeco-Armenian is characterized predominantly by J2+R1b related lineages, while Indo-Iranian by J2+R1a related lineages. The evidence for Tocharian would involve J2+R1b related lineages.  Overall, it would appear that the earliest J2 core of PIE affected two different groups of populations living on complementary sides of the Caspian:

• The trans-Caspian R-M17 population followed an early (3rd, or late 4th millennium BC?) north-west trajectory into Europe (associated with northern European groups such as Balto-Slavic and Germanic) as well as a later expansion (2nd millennium BC? associated with climatic deterioration in BMAC) that brought Iranian speakers to the steppe, as well as to Iran, and Indo-Aryans to South Asia.
• The cis-Caspian, trans-Caucasian R-M269 population followed an early (late 4th millennium, early 3rd millennium?) expansion into Europe, probably together with J2 in the Balkans (Graeco-Phrygian, perhaps Thracian), and arriving in the form of Bell Beakers in Western Europe (Italo-Celtic), as well as a later (2nd-1st millennium BC?) expansion to the east (Tocharians)

This long excursus was necessary as a preamble to an explanation on what happened in Europe itself, which brings us back to the topic of the current paper:

• The lack of structure between regions A and B with respect to haplogroup J, together with the great difference in levels of this haplogroup between Italy and the Celtic world,  suggests that Italian J-related lineages  may have been inflated in proto-historical and historical times. There are candidates a-plenty: Greeks, Etruscans, Trojans to name but three. Excess of J in Italy, relative to the Celtic world, clearly relates to the abundant traditions of eastern origins for the historical groups of Italy.
• It would appear that during proto-history, most of Europe was dominated by three sets of IE people (R-M269 in the west, who had transmitted Proto-Celto-Italic; R-M17 in the northeast of Proto-Balto-Slavic speech, and Proto-Germanic in-between, participating in both worlds, and --appropriately-- often linked with either Italo-Celtic or Balto-Slavic linguistically)
• There were other (now-extinct) groups as well: the Illyrians vs. Thracians in the Balkans with complementary Y chromosome distributions, the former including an extra chunk of aboriginal legacy (haplogroup I), no doubt due to the much more difficult terrain of the western Balkans. These are contrasted with our final group, the Greeks who straddled three worlds (the Paleo-Mediterranean world of the first farmers, the Thraco-Phrygian world linked to the Indo-Iranians at a deeper level, and the Anatolian world)

The boundaries between these various groups were a little blurred in the course of history. But, apparently, they were still a little clearer during the Middle Ages, and probably much clearer before the Völkerwanderung of the Germans, and the expansion of the Slavs.


Geneticists are executing a remarkable pincer movement, zeroing in on the period of European ethnogenesis from both the remote past and the present: through a study of ancient DNA from the dawn of history, they are beginning to understand how Europe was peopled, layer after layer of settlement; and through the study of surnames and toponyms they are drilling ever deeper into the pre-genealogical past. Together with much anticipated technological progress related to full genome sequencing and ancient DNA extraction, it will not be long before the history of Europe will be laid bare in remarkable detail.

PLoS ONE 7(7): e41885. doi:10.1371/journal.pone.0041885

Pasture Names with Romance and Slavic Roots Facilitate Dissection of Y Chromosome Variation in an Exclusively German-Speaking Alpine Region

Harald Niederstatter et al.

The small alpine district of East Tyrol (Austria) has an exceptional demographic history. It was contemporaneously inhabited by members of the Romance, the Slavic and the Germanic language groups for centuries. Since the Late Middle Ages, however, the population of the principally agrarian-oriented area is solely Germanic speaking. Historic facts about East Tyrol's colonization are rare, but spatial density-distribution analysis based on the etymology of place-names has facilitated accurate spatial mapping of the various language groups' former settlement regions. To test for present-day Y chromosome population substructure, molecular genetic data were compared to the information attained by the linguistic analysis of pasture names. The linguistic data were used for subdividing East Tyrol into two regions of former Romance (A) and Slavic (B) settlement. Samples from 270 East Tyrolean men were genotyped for 17 Y-chromosomal microsatellites (Y-STRs) and 27 single nucleotide polymorphisms (Y-SNPs). Analysis of the probands' surnames revealed no evidence for spatial genetic structuring. Also, spatial autocorrelation analysis did not indicate significant correlation between genetic (Y-STR haplotypes) and geographic distance. Haplogroup R-M17 chromosomes, however, were absent in region A, but constituted one of the most frequent haplogroups in region B. The R-M343 (R1b) clade showed a marked and complementary frequency distribution pattern in these two regions. To further test East Tyrol's modern Y-chromosomal landscape for geographic patterning attributable to the early history of settlement in this alpine area, principal coordinates analysis was performed. The Y-STR haplotypes from region A clearly clustered with those of Romance reference populations and the samples from region B matched best with Germanic speaking reference populations. The combined use of onomastic and molecular genetic data revealed and mapped the marked structuring of the distribution of Y chromosomes in an alpine region that has been culturally homogeneous for centuries.

Link

Iceman stories begin arriving!

The National Geographic has info, a teaser for an October 26 Nova special:

The genetic results add both information and intrigue. From his genes, we now know that the Iceman had brown hair and brown eyes and that he was probably lactose intolerant and thus could not digest milk—somewhat ironic, given theories that he was a shepherd. Not surprisingly, he is more related to people living in southern Europe today than to those in North Africa or the Middle East, with close connections to geographically isolated modern populations in Sardinia, Sicily, and the Iberian Peninsula. The DNA analysis also revealed several genetic variants that placed the Iceman at high risk for hardening of the arteries. ("If he hadn't been shot," Zink remarked, "he probably would have died of a heart attack or stroke in ten years.") Perhaps most surprising, researchers found the genetic footprint of bacteria known as Borrelia burgdorferi in his DNA—making the Iceman the earliest known human infected by the bug that causes Lyme disease.

It seems that my prediction that the Iceman will turn out to be Mediterranean in terms of his autosomal genetic components was right!

I don't get, however, how lactose intolerance is incompatible with being a shepherd, since milk is widely used in southern Europe both as a raw product and for its cheese. I lack the lactose tolerance gene myself, but that doesn't keep me from having a glass of milk nearly every day. Lactose tolerance makes it possible for people to drink lots of milk; lactose intolerance does not make it impossible for them to drink any, or to enjoy its secondary products (such as cheese and butter).

Tyrolean Iceman week is upon us! (and some migrationist thoughts)

After a long wait, this week we are likely to read (and for the lucky few, hear) the long-awaited results of the genetic analysis of Ötzi, the Tyrolean Iceman. The talk on "Sequencing the Tyrolean Iceman" is scheduled for October 21 during the MUMMIES FROM THE ICE, 2nd BOLZANO MUMMY CONGRESS.

Thanks to intelligence from one of my readers, we already know that the Iceman belonged to Y-haplogroup G2a4. My experience with similar publicity bonanzas leads me to believe that the announcement at Bolzano may coincide with a major journal publication, but I have no inside information. Well, Friday is the day when the next issue of Science is due, so I'll take a bet on that journal having an Ötzi special this week. But, even if I'm wrong, there are likely to be media stories from the mummy congress itself; we will soon find out more.

(If any kind souls want to tip me, on promise of absolute confidentiality, feel free to do so: my e-mail's at the bottom)

Why is Ötzi so important? We already have genome sequence data on a ~4,000 year old Paleo-Eskimo, and even older Neandertals and Denisovans. Ötzi, at more than 5,000 years, will become the oldest Homo sapiens sequenced so far.

From a place like Europe, we almost never get anything other than bones and teeth from that period. An ice mummy is a real treasure, as it contains non-osseous tissues preserved in a natural refrigerator. Ötzi is likely to yield as good an ancient DNA genome as we are likely to get from prehistoric Europe anytime soon.

And, Ötzi's age is equally important, since he lived during the early Copper Age, at a time when humans in Europe transcended the use of bone and stone in their tool-making, and started using copper. Humans had used metals before (including gold and iron) in a haphazard way, but it was during the Copper and subsequent Bronze Age that there is clear evidence that metalworking began to transform society.

Ötzi's genome will be extremely important for a different reason: for a long time a conflict has simmered in archaeology between idea diffusionists, demic diffusionists, and migrationists.

1. Idea diffusionists aka proponents of acculturation propose that ideas (such as the idea of crop-raising or metal-working) spread without large movements of people. They predict that Europeans did not change much since the Paleolithic, and Neolithic/post-Neolithic processes have little affected them.
2. Demic diffusionists propose that humans behave like mindless automata, random walking across the landscape, mating with whom they find, and filling up a continent by the accretion of millennia-long processes of diffusion. They predict that Europeans are a fairly smooth cline of Neolithic+Paleolithic constituent elements from southeast to northwest.
3. Migrationists adhere to an older and much-maligned arrows-on-the-map paradigm, whereby humans intentionally decide to move from A to B, even across great distances. According to this idea, colonists sometimes mix with/sometimes kill/sometimes avoid pre-existing inhabitants. Migrationists predict that prehistoric Europe was a dynamic patchwork of genetic-cultural units entering the continent from different routes at different times, gradually forming the cornucopia of its proto-historical ethnic groups.

It's been about two years since I came out as a migrationist. In my view, the colonization of Europe was less a random process and more akin to the much later colonization of the Mediterranean and Black Sea by the Greeks, and of the Americas by Europeans. We can envision initial forays of exploration, prompted by either curiosity or tales of strange sights and great riches (be it the riches of Marco Polo's East, El Dorado, the Golden Fleece, etc.). These were followed by colonists, either pushed from their homelands by social/economic malaise, or pulled towards their destinations by opportunity, establishing long-range communication/trade networks. Finally, more people could flow along the established routes in a directional, intentional flow of people.

Most of the ancient DNA published in the last few years has tended to support the migrationist paradigm. Indeed, we are uncovering even weirder data points every time we look. Who would have thought a few years ago, that Australian aborigines would show ties to Siberia, some prehistoric central-eastern Europeans to modern East Asians, and neither Mesolithic nor Neolithic Europeans any clear ties to modern ones?

It seems that the surest bet is on the unexpected, so I am hopeful that the Tyrolean Iceman will have some surprises in store for us; these may upset existing paradigms, but will pave the way for new ones.

Ötzi, the Tyrolean Iceman belonged to Y-haplogroup G2a4

G was the third most popular choice in my recent poll.

We now have G2a3 from Neolithic Linearbandkeramik in Derenburg and G2a in Treilles in addition to Ötzi from the Alps. G2a folk got around. He joins Stalin and Louis XVI as a famous G2a.

It was already clear with the discovery of G2a in France and Central Europe, that this otherwise uncommon present-day haplogroup in Europe was more prominent during the Neolithic, and Ötzi's data point seals the case.

In a sense, the triple G2a finds in Neolithic Europe confirm the origins of the European Neolithic population in West Asia, but renew the mystery as to how all the rest of the "players" of the European Y-DNA scene appeared on the scene, with everything except G and I first appearing in the ancient DNA record after the end of the Neolithic.

Ötzi has been added to the ancient Y-chromosome studies page.

Craniofacial morphology in Austrian Early Bronze Age

J Anthropol Sci. 2011 Jul 1. [Epub ahead of print]

Craniofacial morphology in Austrian Early Bronze Age populations reflects sex-specific migration patterns.

Pellegrini A, Teschler-Nicola M, Bookstein F, Mitteroecker P.

Abstract
The Early Bronze Age (2.300-1.500 BC) in lower Austria consists of three synchronous regional manifestations (Únetice, Unterwölbling, and Wieselburg cultures). The bearers of these cultures inhabited a relatively small geographic area and shared similar ecological conditions, but previous studies revealed population differences in skeletal morphology. We analyzed the cranial morphology of 171 individuals of these populations with a geometric morphometric approach in order to compare different migration scenarios. We find significant mean form differences between populations and between sexes. In a principal component analysis, the Wieselburg population, located southwest of the Danube, largely separates from the Únetice population north of the Danube, whereas the southwestern Unterwölbling group, which played a central role in trading bronze objects, overlaps with both. The Böheimkirchen group, inhabiting the southwestern Danubian area in the later phase of the Early Bronze Age, differs from the chronologically older Unterwölbling group. Geographic distance between six sites and position relative to the river Danube accounted for 64% of form distance variation; the effect of the river Danube was considerably larger than hat of geographic distance per se. As predicted for a patrilocal system in which females have a larger marriage domain than males, we found that female mean forms are more similar to each other than male mean forms. Geographic conditions explained more than twice as much variation in females as in males, suggesting that female migration was more affected by geographical constraints than male migration was.

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Brown-eyed Oetzi the Iceman

Heike Engel-21Lux / Sdtiroler Archologiemuseum / National Geographic Deutschland

Iceman Oetzi gets a new face for 20th anniversary

BOLZANO, Italy — Iceman Oetzi, whose mummified body was famously found frozen in the Italian Alps in 1991, will get a new face for the 20th anniversary of his discovery.

As part of a new exhibit at the South Tyrol Museum of Archaeology in Bolzano (www.iceman.it), two Dutch experts -- Alfons and Adrie Kennis -- have made a new model of the living Oetzi, this time with brown eyes.

Indeed, recent research has shown the Iceman, now approaching the tender age of 5,300 years, did not have blue eyes as previously believed.

The Kennis model was created based on three-dimensional images of the mummy's skeleton as well as the latest forensic technology, and will go on display on March 1 until January 15, 2012.

The Bolzano museum is also planning to improve conservation of the mummy by using pure nitrogen, which should help it keep its relatively young appearance by eliminating oxidation.

Oetzi was discovered on September 19, 1991 by a German couple in a glacier in the Oetztal Alps in South Tyrol, northern Italy.

The Iceman, who is believed to have died around the age of 45, was about 1.60 metres (5 foot, 3 inches) tall and weighed 50 kilogrammes (110 pounds), about average for his time. If he had lived today, he would have worn size 38 shoes.

But Oetzi's notoriety has also been linked to a supposed curse surrounding the mummy, after several people -- authors, researchers, even mountain guides -- who came in contact with it died over the years.

One of the couple who discovered Oetzi was found dead after a mountain hike in 2004. A forensic expert, who had closely examined the find, meanwhile died in a car accident in 1992 on his way to a lecture about Oetzi.

Picture from an Italian story on the same topic