Oxhide ingots in Scandinavian rock art

Antiquity / Volume 89 / Issue 343 / February 2015, pp 191-209

Representations of oxhide ingots in Scandinavian rock art: the sketchbook of a Bronze Age traveller?

Johan Ling and Zofia Stos-Gale

Bronze Age trade networks across Europe and the Mediterranean are well documented; Baltic amber and bronze metalwork were particularly valued commodities. Here it is argued that demand for copper and tin led to changes in Scandinavian trade routes around 1600 BC, which can be linked to the appearance of figurative rock art images in southern Scandinavia. Images identified as oxhide ingots have been discovered in Sweden and suggest that people from Scandinavia were familiar with this characteristically Mediterranean trading commodity. Using trace element and lead isotope analysis, the authors argue that some bronze tools excavated in Sweden could have been made of Cypriot copper; these two discoveries suggest that Scandinavians were travelling to the Mediterranean, rather than acting through a middle man.

Link

More ancient Scandinavians (Skoglund, Malmström et al. 2014)

A new paper has just appeared in Science which adds new ancient Swedish hunter-gatherer samples, as well as a new Gökhem2 Swedish farmer. Much lower quality data from the same archaeological sites were studied in 2012 by much the same team, but the new study has sequenced several new Pitted Ware individuals from Ajvide, as well as a  Mesolithic Swede.

The Swedish hunter-gatherers appear to be similar to those of Lazaridis et al. (2013) in that their ancestry is a mixture of both European hunter-gatherers like LaBrana1 and ~15% of something related to MA1, which seems quite close to the 19% of ANE ancestry for the older Motala hunter-gatherer also from Sweden. The finding of Y-haplogroup I2a1 also parallels the Motala hunter-gatherers, so everything seems quite consistent with the Mesolithic Swedes being genetically very close to the Pitted Ware Neolithic ones. However, there is one difference in that the new hunter-gatherers were ancestral for SLC24A5 while the Motala one was derived (this is the "skin lightening" allele that was curiously missing in both Iberia and Luxembourg hunter-gatherers).

The authors also find that the Iceman and Gökhem2 are a mixture of Basal Eurasians and something related to hunter-gatherers. A interesting new detail is that the Swedish farmer had more of the hunter-gatherer ancestry than the Iceman (the estimated difference in their non-Basal Eurasian ancestry is 77.2-56=21.2%) which seems reasonably close to the 16% difference in the related "Atlantic_Baltic" ancestry for the previous lower-quality Gok4 farmer and the Iceman I estimated in 2012.

Finally, the authors also study the genetic diversity of the Swedish hunter-gatherers:

The Scandinavian Neolithic hunter-gatherer group had significantly lower conditional nucleotide diversity (0.181±0.0015) compared to the Scandinavian Neolithic farmer group (0.201±0.0038, Figs 3A and S9). While the specific properties of ancient DNA may still affect comparisons with sequence data from modern-day individuals, the conditional nucleotide diversity in the hunter-gatherers was also lower than in any modern-day European and a Chinese population (22) analyzed using the same approach as for the ancient groups.

It is not easy to estimate nucleotide diversity with low coverage data (because you can't tell whether a sample is heterozygous in some position if you only have a handful of reads covering it), but the authors cleverly use the fact that they have multiple individuals from the hunter-gatherer population to estimate this. The low diversity in hunter-gatherers also parallels the finding of low genetic diversity in the Luxembourgeois Mesolithic hunter-gatherer, so it does seem that hunter-gatherers in Europe were a very low diversity population, which seems reasonable for people engaging in foraging (which does not allow for growth to large population numbers) and having ancestors who endured the Ice Age in Europe.

The last few months have been extremely generous in new ancient DNA studies and I hope that more stuff is coming this year as in 2013.

UPDATE: Also important (from the Independent):

“We see clear evidence that people from hunter-gatherer groups were incorporated into farming groups as they expanded across Europe. This might be clues towards something that happens also when agriculture spread to other parts of the world,” Dr Skoglund said.

Science DOI: 10.1126/science.1253448

Genomic Diversity and Admixture Differs for Stone-Age Scandinavian Foragers and Farmers

Pontus Skoglund, Helena Malmström et al.

Prehistoric population structure associated with the transition to an agricultural lifestyle in Europe remains contentious. Population-genomic data from eleven Scandinavian Stone-Age human remains suggest that hunter-gatherers had lower genetic diversity than farmers. Despite their close geographical proximity, the genetic differentiation between the two Stone-Age groups was greater than that observed among extant European populations. Additionally, the Scandinavian Neolithic farmers exhibited a greater degree of hunter-gatherer-related admixture than that of the Tyrolean Iceman, who also originated from a farming context. In contrast, Scandinavian hunter-gatherers displayed no significant evidence of introgression from farmers. Our findings suggest that Stone-Age foraging groups were historically in low numbers, likely due to oscillating living conditions or restricted carrying-capacity, and that they were partially incorporated into expanding farming groups.

Link

Origin of copper ores of Nordic Bronze Age

Journal of Archaeological Science doi:10.1016/j.jas.2013.07.018

Moving metals II: provenancing Scandinavian Bronze Age artefacts by lead isotope and elemental analyses 

Johan Ling et al.

The first part of this research published previously proved without doubt that the metals dated to the Nordic Bronze Age found in Sweden were not smelted from the local copper ores. In this second part we present a detailed interpretation of these analytical data with the aim to identify the ore sources from which these metals originated. The interpretation of lead isotope and chemical data of 71 Swedish Bronze Age metals is based on the direct comparisons between the lead isotope data and geochemistry of ore deposits that are known to have produced copper in the Bronze Age. The presented interpretations of chemical and lead isotope analyses of Swedish metals dated to the Nordic Bronze Age are surprising and bring some information not known from previous work. Apart from a steady supply of copper from the Alpine ores in the North Tyrol, the main sources of copper seem to be ores from the Iberian Peninsula and Sardinia. Thus from the results presented here a new complex picture emerges of possible connectivities and flows in the Bronze Age between Scandinavia and Europe .

Link

The Nordic razor

I had posted a few studies suggesting links between Mycenaean Greece and Scandinavia, and here is another one. From the paper, this ties a bit to my ideas about the establishment of long-range networks associated with metallurgy in the Bronze Age:

It can be seen that there were two, chronologically separate, lines of introduction or transfer of the razor idea from the eastern Mediterranean to northern Europe. The spread of the two-edged razor to Central and Western Europe including Britain and Ireland took place just before or around 1500 BC. The one-edged razor arrived in Scandinavia in the decades before 1400 BC. The two ‘time-slots’ of transfer from the Mediterranean of two types of razors indicate the use of specific long distance networks that were probably in existence beforehand.

Antiquity Volume: 87 Number: 336 Page: 461–472

The Nordic razor and the Mycenaean lifestyle

Flemming Kaul

*Danish Prehistory, The National Museum of Denmark, Frederiksholms Kanal 12, Copenhagen DK 1220, Denmark (Email: flemming.kaul@natmus.dk)

The bronze razor with the horse-head handle appeared in Scandinavia in the fifteenth century BC. Where did it come from and what did it mean? The author shows that the razor had some antecedents in the Aegean, although none of these objects were imported to the north. He argues that the Scandinavian warrior class consciously adopted elements of the Mycenaean warrior package, including a clean-shaven face. This vividly exposes new aspects of the busy and subtle nature of international communication in the Bronze Age.

Link

Links between Mycenaeans and Scandinavia

Three papers on a similar theme. An excerpt from a source mentioned in the second paper:

Det visar sig att alla undersökta svenska föremål utom ett enda - en slaggbit - kommer från gruvor och malmfyndigheter från platser på Cypern, Sardinien, Iberiska halvön, Massif Central i nuvarande Frankrike, Tyrolen samt Brittiska öarna. Kopparn har transporterats hit och i utbyte har man skeppat tillbaka stora mängder bärnsten. Fram träder en bild av en tid då internationella kontakter över stora vatten var självklarheter, och det redan cirka 2000 år innan vikingarna gav sig iväg på sina färder. [Google Translate]: It turns out that all examined Swedish subject except one - a slaggbit - comes from mines and ore deposits from sites in Cyprus, Sardinia, the Iberian Peninsula, the Massif Central in the current France, Tyrol and the British Isles. Copper has been transported, and in return it has been shipped back large amounts of amber. What emerges is a picture of a time when international contacts over large water was obvious, and there are already some 2000 years before the Vikings set off on their journeys.

From the third paper:

Both the lead isotope and chemical analyses have undoubtedly showed that the copper from the 33 Scandinavian Bronze Age artefacts diverges significantly from Scandinavian copper ores and that the copper must have been imported from elsewhere. The results furthermore indicate that there are variations in metal supply that are related to chronology, in resemblance with artefacts from Scandinavia as well as from other parts of Europe indicating analogous trade routes for copper, during the respective periods. Maritime networks and changing sources of metal seem to have been a key feature for Scandinavia in the Bronze Age.

Archaeology, Ethnology and Anthropology of Eurasia

Volume 40, Issue 2, June 2012, Pages 99–103

Grave Circle B at Mycenae in the Context of Links Between the Eastern Mediterranean and Scandinavia in the Bronze Age

I.B. Gubanov

Artifacts from royal burial graves Gamma and Omicron of grave circle B at Mycenae attest to cultural ties between the Eastern Mediterranean elite and that of the Scandinavian Early Bronze Age (mid- and late 2nd millennium BC). The appearance of the running spiral motif and representations of ships with rams in Scandinavia coincide with the beginning of the Mycenaean civilization. These facts, along with the finds of Baltic amber only in the royal burials at Mycenae but not in Crete, suggest that a principal role in the introduction of these cultural elements in Scandinavia during the Scandinavian Bronze Age (periods I–III according to Montelius) was played by the Mycenaean elite.

Link

Journal of Geography and Geology Vol 5, No 1 (2013)

The Bronze Age in SE Sweden Evidence of Long-Distance Travel and Advanced Sun Cult 

Nils-Axel Mörner, Bob G. Lind

The Bronze Age of Scandinavia (1750-500 BC) is characterized by the sudden appearance of bronze objects in Scandinavia, the sudden mass appearance of amber in Mycenaean graves, and the beginning of bedrock carvings of huge ships. We take this to indicate that people from the east Mediterranean arrived to Sweden on big ships over the Atlantic, carrying bronze objects from the south, which they traded for amber occurring in SE Sweden in the Ravlunda-Vitemölla–Kivik area. Those visitors left strong cultural imprints as recorded by pictures and objects found in SE Sweden. This seems to indicate that the visits had grown to the establishment of a trading centre. The Bronze Age of Österlen (the SE part of Sweden) is also characterized by a strong Sun cult recorded by stone monuments built to record the annual motions of the Sun, and rock carvings that exhibit strict alignments to the annual motions of the Sun. Ales Stones, dated at about 800 BC, is a remarkable monument in the form of a 67 m long stone-ship. It records the four main solar turning points of the year, the 12 months of the year, each month covering 30 days, except for month 7 which had 35 days (making a full year of 365 days), and the time of the day at 16 points representing 1.5 hour. Ales Stones are built after the same basic geometry as Stonehenge in England.

Link

Journal of Archaeological Science
Volume 40, Issue 1, January 2013, Pages 291–304

Moving metals or indigenous mining? Provenancing Scandinavian Bronze Age artefacts by lead isotopes and trace elements

Johan Ling et al.

The aim of this study is to further the discussion as to whether copper was extracted locally or imported to Sweden during the Bronze Age or if both of these practices could have coexisted. For this purpose, we have carried out lead isotope and chemical analyses of 33 bronze items, dated between 1600BC and 700BC. Among these are the famous Fröslunda shields and the large scrap hoard from Bräckan and other items from three regions in southern Sweden which are also renowned for their richness in copper ores. It is obvious from a comparison that the element and lead isotope compositions of the studied bronze items diverge greatly from those of spatially associated copper ores. Nor is there any good resemblance with other ores from Scandinavia, and it is concluded that the copper in these items must have been imported from elsewhere. The results furthermore indicate that there are variations in metal supply that are related to chronology, in agreement with other artefacts from Scandinavia as well as from other parts of Europe. Altogether these circumstances open up for a discussion regarding Scandinavia’s role in the maritime networks during the Bronze Age.

Link

Genomic history of Denmark

An announcement from the GeoGenetics centre:

The Genomic History of Denmark.

The centre has received a 36 mill. DKK grant from Univ. of Copenhagen's dedicated 2016-program. Researchers from GeoGenetics in close cooperation with collegues from the National Museum of Denmark and institutes at the University of Copenhagen will make Denmark the first country in the world to map its evolutionary, demographic and health history - from the earliest settlers through to modern times.

DNA and proteins extracted from a Danish collection of archaeological skeletons from the Older Stone Age (5000-3000 BC) will be analysed in order to learn more about the Danish cultural heritage and health history.

Professor and director of the Centre for GeoGenetics Eske Willerslev is project leader.

Skull trauma in Neolithic Scandinavia

I would be interested to know how this Neolithic sample might differ from more recent ones. My limited understanding suggests that between-male violence often has a signalling component whereby an individual's or group's dominance over another is asserted, so the fight often does not go all the way to death, but only until the status quo is manifested by the controlling party or toppled by a challenger.

This type of "signalling" aspect of violent behavior does not apply to male-to-female violence because of the physical strength inequality between the sexes. Indeed, as with violence towards children or the elderly, male-to-female violence may have a "reverse signalling" effect, because it suggests that the perpetrator is unable to fight with "the strong" and is only able to assert physical dominance in "easy fights". On the other hand, such "easy fights" might be more abundant if perpetrators tend to enter fights they can win.

Fight-to-the-death, on the other hand, may occur either by accident (e.g., when the aim is to assert dominance, but the killer underestimates the tolerance of the victim), or by intent (when the aim is physical annihilation, either because reconciliation with the victim is perceived to be impossible, or because the victim's death may help keep other challengers in check).

There may be lots to learn about gender roles and social hierarchy from large palaeoanthropological samples. For example, how much did ideology affect secular patterns of interpersonal violence, and how much did changes in weapon technology (e.g., from Neolithic to Bronze, Iron, and more recently firearms).

Am J Phys Anthropol DOI: 10.1002/ajpa.22192

Patterns of violence-related skull trauma in neolithic southern scandinavia

Linda Fibiger et al.

This article examines evidence for violence as reflected in skull injuries in 378 individuals from Neolithic Denmark and Sweden (3,900–1,700 BC). It is the first large-scale crossregional study of skull trauma in southern Scandinavia, documenting skeletal evidence of violence at a population level. We also investigate the widely assumed hypothesis that Neolithic violence is male-dominated and results in primarily male injuries and fatalities. Considering crude prevalence and prevalence for individual bones of the skull allows for a more comprehensive understanding of interpersonal violence in the region, which is characterized by endemic levels of mostly nonlethal violence that affected both men and women. Crude prevalence for skull trauma reaches 9.4% in the Swedish and 16.9% in the Danish sample, whereas element-based prevalence varies between 6.2% for the right frontal and 0.6% for the left maxilla, with higher figures in the Danish sample. Significantly more males are affected by healed injuries but perimortem injuries affect males and females equally. These results suggest habitual male involvement in nonfatal violence but similar risks for both sexes for sustaining fatal injuries. In the Danish sample, a bias toward front and left-side injuries and right-side injuries in females support this scenario of differential involvement in habitual interpersonal violence, suggesting gendered differences in active engagement in conflict. It highlights the importance of large-scale studies for investigating the scale and context of violence in early agricultural societies, and the existence of varied regional patterns for overall injury prevalence as well as gendered differences in violence-related injuries.

Link

Another look at Neolithic inhabitants of Sweden, using 'euro7'

After my initial analysis of the new Swedish Neolithic data, I decided -like with the Iceman-  to assess them with the euro7 calculator. Check the spreadsheet for comparative values in modern populations.

Of the hunter-gatherers: Ajv70 turns out to be 100% "Northeastern" in this analysis; Ajv52 is 75.1% "Northeastern", 21.8% "Northwestern", 2.9% "Far_Asian", and 0.3% "African". The "Northeastern" component is modal in the Baltic area.

Gok4, the Megalithic farmer was 61.5% "Northwestern", 21.4% "Southeastern", and 17.1% "Southwestern". The "Northwestern" component is modal in Atlantic Europe.

Modern Scandinavians have very little "Southeastern" and "Southwestern" by comparison, and this is probably what accounts for the clear southern origin of Gok4. But, Gok4 does seem to share the major "Northwestern" component with modern Scandinavians.

Two additional 'euro7' components occur in modern Scandinavians, "Caucasus" (~5%) and "Northeastern" (~20-25%). These components also occur in modern populations of the British Isles: the "Caucasus" one at similar or higher frequencies, the "Northeastern" one at lower ones than is the case in Scandinavia.

The fact that these two 'euro7' components are also missing in Oetzi strongly suggests to me that there was a late-Neolithic or post-Neolithic east-to-west migration into Europe from an eastern source area. Given the absence of "Caucasus" in Neolithic Gotland hunter-gatherers, it is a reasonable assumption that this migration may have originated from further south and east, where the "Caucasus" component occurs in modern populations.

There have been several indications linking Northwestern Europe with the Northeast Caucasus region. The latter exhibits high levels of Y-haplogroup R1b, the main Northwest European lineage. It also exhibits unexpectedly high levels of the "Northwest" component. I have little doubt that these twin facts constrain our understanding of the peopling of Western Europe by anchoring it -in some manner- to the Caspian and its environs.

If I had to guess I would propose the following scenario:

• The "Northwestern" component represents the pre-Megalithic first farmers of Northwestern Europe, consisting of Linearbandkeramik farmers emanating from Central Europe and admixing with pre-farming Atlantic hunter-gatherers.
• The "Megalithic" phase of the Neolithic saw the infusion of a new wave of maritime colonists originating in the eastern Mediterranean ("Southeastern") via Iberia ("Southwestern") and reaching their terminus in Scandinavia.
• The last major population movement into Northwestern Europe involved the arrival of a population element from the northern parts of the Near East via the Caucasus, probably originating in the north Iran/Armenia/Azerbaijan/Dagestan "short arc" west and south of the Caspian where there is a local maximum of R1b frequency.

Thankfully, there is ongoing ancient DNA work on both the European steppe and the Balkans/Anatolia, i.e., the two possible conduits through which any additional "players" in the peopling of Europe must have passed through. Together with ancient DNA study of other archaeological cultures in continental Europe itself, (e.g., Corded Ware/Bell Beaker) our picture of prehistoric events is bound to become ever sharper in coming years.And, hopefully, once the actors in the drama are revealed, we can move on to the late Bronze and Iron Ages, to see how they interacted to form the historical peoples of West Eurasia.

A first look at the DNA of Neolithic inhabitants from Sweden

I was eager to get my hands on the ancient DNA from ancient Swedish Neolithic people, as soon as I became aware of it a few months ago. I was finally able to extract SNPs from the data, and I decided to test the samples with my K7b and K12b calculators from the Dodecad Project, as I had also recently done for the Tyrolean Iceman. You can skip to the Results section if you want.

I extracted the following number of SNPs from the different individuals, that were in common with my main HGDP reference:

• 15,734 SNPs: Gok4 (TRB / farmer)
• 15,385 SNPs: Ajv52 (PWC / hunter-gatherer)
• 25,108 SNPs: Ajv70 (PWC / hunter-gatherer)

I did have some trouble with my code for extracting data from the third hunter-gatherer individual, Ire8. This also happens to have the lower number of SNPs as reported by Skoglund et al. so its non-inclusion is probably not a great loss.

K7b and K12b are based on a set of 166,770 SNPs, so I intersected the SNPs of the ancient individuals with them, resulting in:

• 4,054 SNPs: Gok4
• 4,077 SNPs: Ajv52
• 6,631 SNPs: Ajv70

This is, of course, a small number of SNPs, and if we tried to infer structure within West Eurasia with them, we might fail. Nonetheless, by exploiting the structure inferred by the larger number of SNPs in modern populations, and using these to test the ancestry of the ancient ones, we get results that appear to agree fairly well with Skoglund et al. (2012).

A problem with comparing against extant populations, rather than ancestral components is that relationships are averaged: for example, Turks in Skoglund et al. appear quite distant to both Neolithic groups, but we cannot know to what extent this is due to their small levels of central/east Eurasian ancestry and to what extent this is due to their native Anatolian ancestry.

We now have two ancient autosomal DNA sampling locations, and both turned up unexpected results. The Iceman, a Copper Age inhabitant of the Alps resembled modern Sardinians. A Megalithic Swedish farmer resembled Southern Europeans, while his hunter-gatherer contemporaries were outside the range of modern variation. These results should give us caution about the identity of past populations elsewhere. 

As I have argued elsewhere, the past seems to have been much more dynamic than many had suspected, and the people that walked and rowed to the ends of the Earth during the Upper Paleolithic did not suddenly grow fetters or decided to stay put during the Neolithic as many "Paleolithic continuity" adepts had proposed.

A couple more caveats:

1. Irrespective of their actual origin, these individuals would still be inferred to be some admixture of the ancestral components adding up to 100%. This hints at their affiliations, but does not -in itself- supply evidence of their absolute proximity to the ancestral components.
2. As more and more ancient individuals are sampled, we will be able to generate genuine ancient populations that are ancestral to modern ones. When that happens, we can directly test modern individuals against panels of ancient ones. For the time being, we have to make do with the reverse, i.e., test ancient individuals against panels of modern ones.

RESULTS


In all comparisons with other K7b results, you should take into account the much smaller number of SNPs used on the Neolithic remains from Sweden.

The K7b results are below. Consult the spreadsheet for comparative values in modern populations.

The K12b results are below. Consult the spreadsheet for comparative values in modern populations.

The raw percentages can be seen below; I have also added the results previously calculated for Oetzi, the Tyrolean Iceman:

DISCUSSION


The results for the two hunter-gatherer samples are as expected predominantly "Atlantic_Baltic" at K=7 and "North_European" at K=12. Since these two samples are outside modern variation, it is expected that their mapping may have added noise; see this post about the dangers lurking at the edges of variation.

Nonetheless, the results can be interpreted as reflecting the fact that the "Atlantic_Baltic" and "North_European" components partially reflect the pre-farming population of Europe. At K=12 it is noteworthy that there is a minor "Atlantic_Mediterranean" admixture in the two hunter-gatherer samples. In my opinion, this may reflect either some level of admixture with the incoming farmers and/or the pre-farming component (but of Western European rather than Baltic type) that may also exist in these foragers. On balance, however, the "North_European" component far outweighs the "Atlantic_Mediterranean" one, which is also consistent with their location (Gotland) which ties them to the Baltic rather than Atlantic Europe.

The farmer sample is remarkable in that, like the Tyrolean Iceman, she seems to be made up entirely of "Atlantic_Baltic" and "Southern" at K=7. There is a hint that the order of the two components are reversed in Gok4 relative to Oetzi, which probably makes sense. The third major West Eurasian component at this level of resolution, the "West Asian" one is conspicuously absent. This component -bimodal in the Caucasus and Balochistan, and strongly represented in the highlands of West Asia in between the two- does occur at ~10% in modern Northern Europeans, so its absence in all Neolithic samples so far hints at its later introduction into at least parts of Europe.

The result at K=12 is fascinating, since Gok4 turns out to be 81% "Atlantic_Mediterranean", and, like Oetzi, with a noticeable "Southwest_Asian" strain. The "Atlantic_Mediterranean" component is bimodal in modern Sardinians and Basques, while the "Southwest_Asian" radiates from southern parts of the Near East into Mediterranean Europe.

Gok4 appears to be "even more Atlantic_Mediterranean" than any modern population. As I mention in the original post, the inhabitants of megalithic monuments  of North-Western Europe were believed by Coon to belong to a "Long Barrow type" which he considered ancestral to the modern "Atlanto-Mediterranean" type. Here is his description:

Toward the end of the Neolithic period, the western Mediterranean countries were invaded by seafarers of a tall, exceptionally long-headed Mediterranean variety; some of these invaders passed through the Straits of Gibraltar, whence they also invaded the British Isles and Scandinavia. 

I would say that the evidence is not incompatible with this scenario. We must probably wait to see whether Gok4 was descended from seafarers from the eastern Mediterranean (where the "Southern" component is modal), following the Mediterranean and then Atlantic coasts up to Scandinavia, or whether they are descended from a different group of people who followed the plains and river valleys of the Balkans and Central Europe and arrived to the north via the inland route. The strong Atlantic_Mediterranean result, coupled with high levels of allele sharing in Cyprus, Greece, France, and the Netherlands, but not particularly in the northern Balkans leads me to prefer the maritime colonization scenario, at least for now. Ancient DNA from more European regions will hopefully help us better understand "what really happened in prehistory."

Ancient DNA from Neolithic Sweden (Skoglund et al. 2012)

A new paper in Science solidifies the case for migration as the cause for the diffusion of agriculture in Europe. Discontinuity between early Neolithic farmers and Mesolithic foragers in Central Europe had provided strong hints about this discontinuity, and these were confirmed by other ancient European DNA, e.g., from Treilles, or the Tyrolean Iceman. The case now appears irrefutable, that people not ideas were involved in the spread farming to the northern fringes of Europe.

If we were to ever find signs of acculturation, the north-eastern corner of Europe may be best place to look for it. Agriculture arrived late to Scandinavia and the Baltic, so there was maximum opportunity for Neolithic groups in the area to acquire pre-Neolithic genes from acculturated farmers during their ~2ky long journey from the Aegean. Conversely, forager populations persisted here longer than elsewhere in Europe, both due to the remoteness of the area and the relative unsuitability of the Neolithic package brought from more southern latitudes.

During the Neolithic period there still existed foragers in Scandinavia who belonged to the Pitted Ware (PWC) culture. These have been the object of a previous mtDNA study, which found them to be strongly differentiated from contemporaneous Funnel Beaker or Trichterbecherkultur (TRB) farmers. The latter were farmers who were also associated with Megalithic monuments in northern Europe.

A recent article by Rowley-Conwy (2011), from which the figure on the left is taken, gives some archaeological perspective on the Neolithic of southern Scandinavia:

This farming spread must have been by boat. There were no native aurochs on Zealand (Aaris-Sorensen 1980), so the early cattle at Akonge were definitely imported. Farther north, agriculture was probably carried by boat up the coasts, an easier method of travel than overland (see above). Baltic crossings would require longer open-water voyages than in the Cardial or LBK. Irish curraghs can, however, make substantial voyages and weather considerable seas (Hornell 1938, sec. 5:17–21), and a large one has even crossed the Atlantic (Severin 1978).

...

The agricultural arrival in southern Scandinavia thus appears sharp. Gradualist views of Late Mesolithic developments can be discounted despite the spread of shoe-last axes beyond the farming frontier. Western Norway presents a similar pattern: axes and ceramics were in circulation for over a millennium beyond the farming boundary.

This was the dusk of the European foragers: whatever their contribution to subsequent European populations, their way of life would soon give way to that of the farmer and shepherd. The Pitted Ware culture can indeed be seen as their "last stand", the last time in prehistory when they could co-exist on fairly equal terms with their farmer neighbors.

Hence, it is very exciting to be able to study DNA from this place and time directly, as Skoglund et al. do in a new paper which reports the successful extraction and analysis of ancient DNA from 3 PWC hunter-gathers and one TRB farmer of about ~5,000 years ago:

The Neolithic farmer sample ('Gok4') was excavated from a megalithic burial structure in Gokhem parish, Sweden, and has been directly 14C-dated to 4,921 ± 50 calibrated years BP (calBP), similar to the age (5,100-4,900 calBP) of the majority of other finds in the area (15). There were no indications from the burial context suggesting that Gok4 was different from other TRB individuals (15, 16), and strontium isotope analyses indicate that Gok4 was born less than 100 km from the megalithic structure, similar to all other analyzed TRB individuals from the area (17). The three Neolithic hunter-gatherer samples were excavated from burial grounds with single inhumation graves on the island of Gotland, Sweden, for which associated remains have been dated to 5300-4400 calBP (16).

We must keep in mind that a limited amount of DNA sequence was extracted, which corresponds to a few tens of thousands of SNPs in common with the best modern SNP set used; this corresponds to ~5% of the genome, with different success rates for the four sampled individuals. We must also not forget that these are farmers and foragers from a single point in space-time, and from the periphery of Europe, so we should be cautious in generalizing about the Neolithic transition in other parts of Europe.

Nonetheless, the new study reveals two important pieces of information:

First, the 3 PWC individuals are strongly differentiated from the single TRB one:

Regardless of the underlying model, our study provides direct genomic evidence of stratification between Neolithic cultural groups separated by less than 400 km, differentiation which encapsulates the extremes of modern-day Europe, and appears to have been largely intact for ~1,000 years after the arrival of agriculture.

So, it appears that these individuals lived at roughly the same time and within a small area of Europe, and yet they are as different from each other as the most distant current European populations are. These were not simply drawn from the same or similar populations, some of them deciding to take up farming while others to practice fishing and hunting. These were different populations who maintained their distinctiveness long after "first contact".

Two models have dominated European prehistory in recent decades: acculturationists claimed that the Neolithic package of domesticated plants and animals was transmitted across the continent while the people largely stayed put, while demic diffusionists claimed that people did move, but -at least in the most popular version of the model- that they gradually intermarried with local hunter-gatherers, forming a genetic cline of ancestry, at the far end of which the farmers were mostly derived from local foragers.

One could very well say that the acculturationist model views prehistoric people as smart folk with no legs, apparently ready to take up a good new idea, but reluctant to leave their birthplace. The demic diffusionist model, on the other hand, viewed them as mindless automata, moving across the landscape with little purpose, marrying who they met, and filling a continent in much the same way that gas molecules end up filling a room into which they are introduced.

Both these models are now revealed to be wrong: rather, it seems that "leapfrog" colonization may be responsible for the spread of agriculture and its associated technologies (such as Megalithism) across Europe. In this model, farmers lept from place to place across the landscape intentionally, preserving their gene pool and largely ignoring the pre-existing foragers of the landscape.

Of course, farmer and hunter eventually did mix, and hunting cultures became extinct. But, this was a process that seems to have been complete after 4,000 years BP. Acculturation did eventually happen, and agriculturalists did eventually diffuse to every corner of Europe. But, these are events that happened after the initial group(s) of pioneers had opened the frontier. In this respect, the colonization of Europe bears some resemblance to the settlement of the Americas by Europeans: it happened by leaps and bounds, and the early waves of explorers and pioneers may have opened the landscape but did not immediately fill it: this happened later as a result of demographic growth and new waves of migration, with the extant populations being differentially descended -in different proportions- from migrants and natives.

The second important point of the new study is the revelation that the single Neolithic individual from northernmost Europe was similar to extant southern Europeans:

To more closely investigate the genetic similarity of extant European populations (22, 24) to Neolithic humans, we determined for each SNP and each extant population the average frequency of the particular allele found in either the Neolithic hunter-gatherers or the Neolithic farmer (16). The Neolithic hunter-gatherers shared most alleles with Northern Europeans, and the lowest allele sharing was with populations from Southeastern Europe (Fig. 3A). In contrast, the Neolithic farmer shared the greatest fraction of alleles with Southeastern European populations (Cypriots and Greeks), and showed a pattern of decreasing genetic similarity for populations from the Northwest and Northeast extremes of Europe (Fig. 3B). Individuals from Turkey stand out by low levels of allele sharing with both Neolithic groups, possibly due to gene flow from outside of Europe, but all other European populations can roughly be represented as a cline where allele sharing with Neolithic hunter-gatherers is negatively correlated with allele sharing with Neolithic farmers (Fig. 3C). 

Panel C from the allele sharing figure (left) suggests why we should be cautious about trying to reconstruct European prehistory on the basis of a simple 2-way model of admixture between farmers and hunters.

It is true that extant European populations do fall on a clear cline between them that is strongly significant (R=-0.58, p=0.0029). This means is that they are different to each other in the same ways that farmers/hunters were different from each other. But, this still leaves about 2/3 of the variance unexplained: this may be partly due to the "noise" added by the small number of SNPs, and partly by the contribution of other ancestral groups to extant variation. One of these groups may be the east Eurasian element which must contribute to the differentiation of Turks from Europeans. But, there were probably other West Eurasian elements not represented by the two Neolithic groups: the Mesolithic Pitted Ware individuals have been previously assigned predominantly to mtDNA haplogroup U, which forms a minority in extant Europeans; and a handful of Neolithic samples (LBK, Oetzi, Treilles) have failed to turn up any signs of the dominant R1 Y-haplogroup of extant Northern Europeans. There must be other actors to be revealed in the unfolding story of European origins.

A strong hint for this can also be found in the quite unexpectedly low "TRB" allele sharing of groups from the Northwestern Balkans. This is quite unexpected, as the area is widely believed to be a conduit through which agriculture spread into Central Europe. It is also an area with world maxima of Y-haplogroup I, a lineage which may be a remnant of Paleolithic Europeans, and correspondingly low levels of haplogroups that appear to have arrived later into Europe.

Another important point is that levels of allele sharing between these Neolithic individuals and modern Europeans is generally lower than between most pairs of modern European populations. This is, in part, expected, since the Neolithic specimens are separated by modern populations by ~5ky of evolution, but may also be due to the contribution of unsampled groups to the ancestry of the latter.

From the paper:

We found that compared to a worldwide set of 1,638 individuals (21-23), all four Neolithic individuals clustered within European variation (Fig. S5). However, when focusing the analysis on 505 individuals of European and Levantine descent, the three Neolithic hunterg atherers appeared largely outside the distribution of the modern sample, but in the vicinity of Finnish and northern European individuals (Fig. 1A). In contrast, the Neolithic farmer clustered with southern Europeans, but was differentiated from Levantine individuals. This general pattern persisted for a geographically broader reference data set of 1,466 extant individuals of European ancestry (22, 24) (Fig. 1B), for a much larger number of markers from 241 individuals in the 1000 genomes project (25) (Fig. 1C), and using model-based clustering (26, 27) (Fig. 1D). Although all Neolithic individuals were excavated in Sweden, neither the Neolithic farmer nor the Neolithic hunter-gatherers appeared to cluster specifically within Swedish variation, a pattern that remained also for a larger sample of 1,525 individuals from across Sweden (28) (Fig. S9, Fig.S21-22).

I will try to perform an analysis of these 4 Neolithic Europeans, as I did with the Iceman, and see how they relate to a larger number of populations: for example, the Mesolithic hunter-gatherers have the highest allele sharing with Poles: do they share even more with Lithuanians and other Baltic peoples? The Neolithic farmer is by far closer to Cypriots: are there any populations of the Near East that are close to it as well?

Hopefully, in the near future we may get our first glimpses of genuine Mesolithic Europeans:

In our genomic analyses, the Scandinavian Neolithic hunter-gatherers (PWC) have a genetic profile that is not fully represented by any sampled contemporary population (Fig. 1), and may thus constitute a gene pool that is no longer intact or that no longer exists. While the origin of the Neolithic hunter-gatherers is contentious, the similar mtDNA haplogroup composition of PWC individuals (8) (Table 1) and Mesolithic- and Paleolithic individuals (7, 29) indicate some continuity with earlier European populations, but resolving this hypothesis will require pre-Neolithic genomic data.

The continuity between Mesolithic and Neolithic hunter-gatherer populations in the Baltic is supported by craniometric analysis from a recent paper (left), but it is definitely worth investigating whether -despite their strong differentiation- the Neolithic farmers and foragers of Sweden may not have already started -at least partially- the process of amalgamation.

Hopefully we can soon extract more DNA from other Neolithic Europeans, as well as pre-contact European foragers. It is probably in the Copper and Bronze Ages that we are to encounter some the remaining players that formed the European genetic landscape and witness how they all combined to form the proto-historical and recent Europeans.

A Postscript:

Until recently, it had become commonplace in archaeology to seek local origins for most archaeological phenomena. Three years ago, I pointed out that new evidence was pointing towards a major migrationism comeback in our understanding of European prehistory. So, it is worth reviewing what was once thought about the people buried in Swedish megalithic monuments. From Carleton Coon's The Races of Europe, (1936) Chapter IV):

In Sweden, out of twenty-four male crania found in passage graves, only one was brachycephalic; for the most part a pure Long Barrow type is represented. (Section 12) 

The Megalithic Long Barrow people must have come by sea, and they probably came from somewhere in the Mediterranean. (Section 10)

The paper is also discussed in the weekly Science podcast. The supplementary materials are freely available.


Science 27 April 2012: Vol. 336 no. 6080 pp. 466-469 DOI: 10.1126/science.1216304

Origins and Genetic Legacy of Neolithic Farmers and Hunter-Gatherers in Europe

Pontus Skoglund1,*, Helena Malmström1, Maanasa Raghavan2, Jan Storå3, Per Hall4, Eske Willerslev2, M. Thomas P. Gilbert2, Anders Götherström1,5,*,†, Mattias Jakobsson

The farming way of life originated in the Near East some 11,000 years ago and had reached most of the European continent 5000 years later. However, the impact of the agricultural revolution on demography and patterns of genomic variation in Europe remains unknown. We obtained 249 million base pairs of genomic DNA from ~5000-year-old remains of three hunter-gatherers and one farmer excavated in Scandinavia and find that the farmer is genetically most similar to extant southern Europeans, contrasting sharply to the hunter-gatherers, whose distinct genetic signature is most similar to that of extant northern Europeans. Our results suggest that migration from southern Europe catalyzed the spread of agriculture and that admixture in the wake of this expansion eventually shaped the genomic landscape of modern-day Europe.

Link

Report on the symposium on Modern Human Genetic Variation

Joshua Akey summarizes the talks of a recent symposium at the Swedish Royal Academy of Sciences. Two bits of information stand out from his report. The first:

In another talk focused on demography, Mattias Jakobsson (Uppsala University, Sweden) presented novel data on the impact of the agricultural revolution on the genetics of contemporary European populations. Specifically, Jakobsson and colleagues obtained nearly 250 Mb of sequence from three 5,000-year-old remains of Neolithic hunter-gatherers and one Neolithic farmer excavated in Scandinavia. Analysis of these sequences in the context of the present day European gene pool suggests that the spread of agriculture involved the northward migrations of farmers. Thus, these data provide the most direct and compelling support for the demic diffusion model of agriculture (as opposed to cultural diffusion) described to date. 

It seems I have my answer to the what's next question. Jakobsson has been doing some interesting work on the demography of human emergence and dispersal, so it will be interesting to see not only the novel sequences from these Neolithic Scandinavians, but also how they fit into existing models of demic diffusion.

The second bit of information:

Similarly, Jeff Wall (University of California San Francisco, USA) described a novel method for inferring archaic admixture, which he applied to publicly available whole-genome sequence data generated by Complete Genomics. Provocatively, he finds higher rates of introgression in Asians compared to Europeans. An advantage of Wall’s method is that it does not require an archaic genome to infer introgression, and thus he was able to also test the hypothesis that contemporary African genomes have signatures of gene flow with archaic human ancestors. Strikingly, Wall indeed did find evidence of archaic admixture in African genomes, suggesting that modest amounts of gene flow were widespread throughout time and space during the evolution of anatomically modern humans.

I guess that I shouldn't throw explanation #1 out the window yet. Wall was involved in the recent paper on archaic African admixture, which only looked at a small subset of the genome, so it is nice to see that he is now working with full genomes, and that the race to data mine complete genomes for archaic admixture is afoot.

The book of abstracts is online at the symposium site. The Jakobsson paper does seem to agree with our emerging picture of a non-local origin of northern European farmers as well as greater survival of pre-farming populations in the northern periphery of Europe, but it will be interesting to see where exactly extant populations fall on the farmer-hunter/gatherer continuum.

Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Northern Europe 

Mattias Jakobsson
Department of Evolutionary Biology, Evolutionary Biology Centre (EBC), Uppsala University, Sweden 

The prehistoric spread of farming in Europe has garnered intense interest for almost a century, and was one of the first questions to which population genetic data was used to investigate demographic hypotheses. However, the impact of the agricultural revolution on the European gene pool remains largely unknown. We obtained 249 million base pairs of quality-filtered human autosomal sequence data from some 5,000 year-old remains of three Neolithic hunter-gatherers and one Neolithic farmer excavated in Scandinavia, the northernmost fringe of agricultural practice at the time. Applying novel methods to study population structure based on low genome-coverage data, we find that Northern European Neolithic farmers are most similar to modern-day southern Europeans, contrasting sharply to Neolithic hunter-gatherers who are most similar to extant individuals from northern Europe. With most extant European populations appearing genetically intermediate between the two Neolithic groups, our results suggest that migration from the south by a genetically distinct group of humans accompanied the spread of agriculture to geographic regions where hunting and gathering was the mode of subsistence, but that admixture eventually shaped modern-day patterns of genomic variation.

Archaic admixture in the human genome 

Jeff D Wall
Department of Epidemiology & Biostatistics, University of California, San Francisco, USA 

We describe a method that uses patterns of linkage disequilibrium in extant human populations to identify regions of the genome that were inherited from ‘archaic’ human ancestors, such as Neandertals, Homo erectus or H. floresiensis. We validate this approach using two recently published archaic human genomes, and show that several ancient admixture events must have occurred, both within and outside of Africa. We also explore differences in the amount of archaic admixture across different contemporary human populations.

Finally, here is the meeting report:

Investigative Genetics 2012, 3:7 doi:10.1186/2041-2223-3-7

Understanding human evolutionary history: a meeting report of the Swedish Royal Academy of Sciences symposium of modern human genetic variation 

Joshua M Akey

Link (pdf)

Reconstruction of 7,500 years old forager from Norway

Source (in Norwegian)

FOTO: ARKEOLOGISK MUSEUM/UNIVERSITETET I STAVANGER

Swedish population structure (Salmela et al. 2011)

From the paper:

In our earlier study [4], we saw that North European populations exhibited differing amounts of IBS similarity to East Asians so that Finns, especially Eastern Finns, were the most similar. Now we have observed the same phenomenon - though in a smaller degree - within Sweden, where Norrland showed the most of East Asian similarity and Götaland the least. This is consistent with earlier Y-chromosomal studies [13]. In strong contrast, however, neither Norrland nor Eastern Finns showed any increase in similarity to the Vologda Russians, and a similar lack of affinity between Finns and Russians can also be seen in separate datasets [6], [13]. Thus, if the current references are representative of Russians in this respect, the observed affinity to Eastern Asia would not be mediated by contacts with Russians but could reflect an ancient eastern influence predating the arrival of Slavic populations to Northeastern Europe in the end of the first millennium [23]. It remains unclear whether the eastern affinity observed in Sweden would date back to the same era, or rather reflect the amount of later Finnish contacts to the area.

PLoS ONE 6(2): e16747. doi:10.1371/journal.pone.0016747

Swedish Population Substructure Revealed by Genome-Wide Single Nucleotide Polymorphism Data

Elina Salmela et al.

Abstract

The use of genome-wide single nucleotide polymorphism (SNP) data has recently proven useful in the study of human population structure. We have studied the internal genetic structure of the Swedish population using more than 350,000 SNPs from 1525 Swedes from all over the country genotyped on the Illumina HumanHap550 array. We have also compared them to 3212 worldwide reference samples, including Finns, northern Germans, British and Russians, based on the more than 29,000 SNPs that overlap between the Illumina and Affymetrix 250K Sty arrays. The Swedes - especially southern Swedes - were genetically close to the Germans and British, while their genetic distance to Finns was substantially longer. The overall structure within Sweden appeared clinal, and the substructure in the southern and middle parts was subtle. In contrast, the northern part of Sweden, Norrland, exhibited pronounced genetic differences both within the area and relative to the rest of the country. These distinctive genetic features of Norrland probably result mainly from isolation by distance and genetic drift caused by low population density. The internal structure within Sweden (FST = 0.0005 between provinces) was stronger than that in many Central European populations, although smaller than what has been observed for instance in Finland; importantly, it is of the magnitude that may hamper association studies with a moderate number of markers if cases and controls are not properly matched geographically. Overall, our results underline the potential of genome-wide data in analyzing substructure in populations that might otherwise appear relatively homogeneous, such as the Swedes.

Link

Breast-cancer causing mutation in Ashkenazi Jews came from Europeans

The interesting question is: why did this become frequent in AJ? If this mutation did indeed enter the AJ gene pool half a millennium ago, then it may be within the reach of genealogists and historians to uncover its origins.

European Journal of Human Genetics , (1 December 2010) | doi:10.1038/ejhg.2010.203

On the origin and diffusion of BRCA1 c.5266dupC (5382insC) in European populations

The BRCA1 mutation c.5266dupC was originally described as a founder mutation in the Ashkenazi Jewish (AJ) population. However, this mutation is also present at appreciable frequency in several European countries, which raises intriguing questions about the origins of the mutation. We genotyped 245 carrier families from 14 different population groups (Russian, Latvian, Ukrainian, Czech, Slovak, Polish, Danish, Dutch, French, German, Italian, Greek, Brazilian and AJ) for seven microsatellite markers and confirmed that all mutation carriers share a common haplotype from a single founder individual. Using a maximum likelihood method that allows for both recombination and mutational events of marker loci, we estimated that the mutation arose some 1800 years ago in either Scandinavia or what is now northern Russia and subsequently spread to the various populations we genotyped during the following centuries, including the AJ population. Age estimates and the molecular evolution profile of the most common linked haplotype in the carrier populations studied further suggest that c.5266dupC likely entered the AJ gene pool in Poland approximately 400–500 years ago. Our results illustrate that (1) BRCA1 c.5266dupC originated from a single common ancestor and was a common European mutation long before becoming an AJ founder mutation and (2) the mutation is likely present in many additional European countries where genetic screening of BRCA1 may not yet be common practice.

Link

Modern Scandinavians descended (maybe) from Neolithic TRB but not Mesolithic Pitted Ware ancestors

Coming shortly after Bramanti et al. (2009) which discovered a discontinuity between Neolithic farmers from Central and Eastern Europe and the pre-existing hunter-gatherers, a new study examines ancient DNA from northern European populations, extending the picture of discontinuity all the way to Scandinavia. This is one more nail in the coffin of the cultural diffusion hypothesis, and in favor for a demic diffusion of agriculture all the way to the northernmost reaches of Europe.

More on this after I read the full paper.

UPDATE (Sep 25):

From the paper:

Although the hunter-gatherers of Denmark and southern Sweden adopted pottery early on, the Neolithization first took real shape with the appearance of the Funnel Beaker Cultural complex (FBC, also known as the Trichterbecher Kultur [TRB]) some 6,000 years BP (the oldest evidence possible dating back some 6,200 years BP [9]). Atthis time domestic cattle and sheep, cereal cultivation, and the characteristic TRB pottery were introduced into most of Denmark and southern parts of Sweden [6]. Nevertheless,the Neolithization process was slow in Scandinavia, and large are as remained populated by hunter-gatherer groups until the end of the 5th millennium BP.

One of these last hunter-gatherer complexes was the Pitted Ware culture (PWC), which can be identified by its single-inhumation graves distributed over the coastal areas of Sweden and the Baltic Sea islands that lie closest to the Swedish coast. Intriguingly, the PWC first appears in the archaeological record of Scandinavia after the arrival of the TRB (some 5,300 yearsBP) and existed in parallel with farmers for more than a millennium before vanishing about 4,000 years BP (Figure 1).

The authors sampled 3 TRB individuals from "one passage tomb, Gokhem, dated to 5,500–4,500 years BP" which were found to belong to haplogroups H, J, and T, and 19 PWC individuals "from three different sites on the Baltic island of Gotland dated to 4,800–4,000 years BP" which were found to belong to haplogroups J, T, V (one each), "Other" (two), U5 and U5a (three each), and U4/H1b (eight samples).

From the paper:

Given our results, it remains possible that the PWC represent remnants of a larger northern European Mesolithic hunter gather complex. However, it appears unlikely that population continuity exists between the PWC and contemporary Scandinavians or Saami. Thus, our findings are in agreement with archaeological theories suggesting Neolithic or post-Neolithic population introgression or replacement in Scandinavia. To what extent this holds true for other parts of Europe requires further direct testing, although morphological [24, 25], ancient [26], and modern [4, 5] genetic data suggest that this is probably the case.

The results indicate that the PWC was dominated by haplogroup U (about three quarters of the mtDNA gene pool). The inability to resolve between U4 and H1b is due to the portion of the mtDNA sampled. Given (i) the absence of other H subgroups in the large sample, (ii) the higher frequency of U4 in modern populations, (iii) the presence of U4 but not H1b in other pre-farming populations of Europe (after Bramanti et al.), (iv) the absence of U4 in Neolithic populations, (v) the higher coalescence age of U4 compared to H1b, suggesting a deeper ancestry, I am inclined to think that most, if not all of the U4/H1b is actually just U4.

UPDATE II:

Fst between the PWC and modern populations ranged between 0.036 (Latvians) and Saami (0.25). For Swedes and Norwegians they were 0.051 and 0.061. A few conclusions can be drawn from this:

1. The notion of Saami as unmixed descendants of pre-farming Europeans is debunked.
2. Latvians and other populations of the eastern Baltic are the closest (although by no means very close) to the PWC.
3. Swedes and Norwegians are somewhat closer to the pre-farming inhabitants than is the case for Central Europe where Fst=0.086 was estimated by Bramanti et al. (2009)

Traditional physical anthropology held that there were three main elements in northern Europe, which have been given different names, but can be summarized as follows:

1. Narrow- and high-faced populations, a new element in the region, similar to that of Central Europe
2. Broad-faced massive Proto-Europoid populations, the aboriginal inhabitants of northern and eastern Europe
3. Flat-nosed populations with eastern affiliations

To quote Raisa Denisova:

Latvia's most ancient inhabitants tended to be large in size, with large skulls, a distinctly oblong head shape, a broad, high face and a distinctly protruding nose (Denisova 1975). Looking at this data in the context of synchronous populations elsewhere in Europe, we can find specific geographic differentials. This is especially true of the facial width of residents, a factor which has great weight in the specification of race (Denisova 1978). Differences in facial width in Europe became particularly distinctive at the beginning of the Atlantic period, when farming was begun in Europe. At this time, facial width distinctly separated morphological forms in Northern Europe from those in the Mediterranean region -- two distinct geographic regions. Massive, broad-faced morphological forms dominated in northern and northeastern Europe, while gracile, narrow-faced forms are found most often in Middle Europe and the continent's southeastern reaches. During the Atlantic period, narrow-faced populations gradually moved in the northerly and northeasterly direction. They reached the Baltic region only during the Bronze Age. For this reason, during the Mesolithic and Neolithic period, people in the Baltic region (and surrounding regions) had broad faces, a fact which affirms their links to the late Paleolithic populations of Europe.

Modern Scandinavians are more (1) than (2), while modern Balts are more (2) than (1). The mtDNA picture seems fairly consistent with a greater persistence of Proto-Europoid elements among the Balts.

A related public release:

Scandinavians are descended from Stone Age immigrants

Today's Scandinavians are not descended from the people who came to Scandinavia at the conclusion of the last ice age but, apparently, from a population that arrived later, concurrently with the introduction of agriculture. This is one conclusion of a new study straddling the borderline between genetics and archaeology, which involved Swedish researchers and which has now been published in the journal Current Biology.

"The hunter-gatherers who inhabited Scandinavia more than 4,000 years ago had a different gene pool than ours," explains Anders Götherström of the Department of Evolutionary Biology at Uppsala University, who headed the project together with Eske Willerslev of the Centre for GeoGenetics at the University of Copenhagen.

The study, a collaboration among research groups in Sweden, Denmark and the UK, involved using DNA from Stone Age remains to investigate whether the practices of cultivating crops and keeping livestock were spread by immigrants or represented innovations on the part of hunter-gatherers.

"Obtaining reliable results from DNA from such ancient human remains involves very complicated work," says Helena Malmström of the Department of Evolutionary Biology at Uppsala University.

She carried out the initial DNA sequencings of Stone Age material three years ago. Significant time was then required for researchers to confirm that the material really was thousands of years old.

"This is a classic issue within archaeology," says Petra Molnar at the Osteoarchaeological Research Laboratory at Stockholm University. "Our findings show that today's Scandinavians are not the direct descendants of the hunter-gatherers who lived in the region during the Stone Age. This entails the conclusion that some form of migration to Scandinavia took place, probably at the onset of the agricultural Stone Age. The extent of this migration is as of yet impossible to determine."

Related:

• Ancient mtDNA from Iceland
• Roman Age, Viking Age, and Eskomo mtDNA
• Ancient Viking mtDNA from Denmark
• Ancient British mtDNA
• Y chromosomes and mtDNA from Eulau
• Central European farmers not descended from local hunter-gatherers

Current Biology doi:10.1016/j.cub.2009.09.017

Ancient DNA Reveals Lack of Continuity between Neolithic Hunter-Gatherers and Contemporary Scandinavians

Helena Malmström et al.

Abstract

The driving force behind the transition from a foraging to a farming lifestyle in prehistoric Europe (Neolithization) has been debated for more than a century [1], [2] and [3]. Of particular interest is whether population replacement or cultural exchange was responsible [3], [4] and [5]. Scandinavia holds a unique place in this debate, for it maintained one of the last major hunter-gatherer complexes in Neolithic Europe, the Pitted Ware culture [6]. Intriguingly, these late hunter-gatherers existed in parallel to early farmers for more than a millennium before they vanished some 4,000 years ago [7] and [8]. The prolonged coexistence of the two cultures in Scandinavia has been cited as an argument against population replacement between the Mesolithic and the present [7] and [8]. Through analysis of DNA extracted from ancient Scandinavian human remains, we show that people of the Pitted Ware culture were not the direct ancestors of modern Scandinavians (including the Saami people of northern Scandinavia) but are more closely related to contemporary populations of the eastern Baltic region. Our findings support hypotheses arising from archaeological analyses that propose a Neolithic or post-Neolithic population replacement in Scandinavia [7]. Furthermore, our data are consistent with the view that the eastern Baltic represents a genetic refugia for some of the European hunter-gatherer populations.

Link

Genes of Finns revisited

This paper interprets the discrepancy between Y-chromosome and mtDNA results in Finland as the signature of Scandinavian gene flow into the western parts of the country, with the Y-chromosome gene pool of the east (typified by haplogroup N3) preserving the original inhabitants starting from the Holocene deglaciation.

It it is not at all clear, however "who got there first", and as far as I can see, the evidence just tells us there is a substantial east-west difference in Y-chromosomes in Finland, it doesn't really tell us which of the two elements represents the most ancient stratum.

In my opinion, the Finnish gene pool may contain traces of the aboriginal inhabitants, as well as the later eastern elements which brought the Finnish language, and the later still influences by Germanic Scandinavians. Hopefully the northern cold has been generous with DNA preservation and we may get some direct glimpses into the country's genetic history.

Some related posts:

• Genomic substructure in Finns
• Migrations in the Baltic region inferred from Y chromosomes and mtDNA
• Y chromosome variation of Finns

European Journal of Human Genetics doi:

Genetic markers and population history: Finland revisited

Jukka U Palo et al.

Abstract

The Finnish population in Northern Europe has been a target of extensive genetic studies during the last decades. The population is considered as a homogeneous isolate, well suited for gene mapping studies because of its reduced diversity and homogeneity. However, several studies have shown substantial differences between the eastern and western parts of the country, especially in the male-mediated Y chromosome. This divergence is evident in non-neutral genetic variation also and it is usually explained to stem from founder effects occurring in the settlement of eastern Finland as late as in the 16th century. Here, we have reassessed this population historical scenario using Y-chromosomal, mitochondrial and autosomal markers and geographical sampling covering entire Finland. The obtained results suggest substantial Scandinavian gene flow into south-western, but not into the eastern, Finland. Male-biased Scandinavian gene flow into the south-western parts of the country would plausibly explain the large inter-regional differences observed in the Y-chromosome, and the relative homogeneity in the mitochondrial and autosomal data. On the basis of these results, we suggest that the expression of 'Finnish Disease Heritage' illnesses, more common in the eastern/north-eastern Finland, stems from long-term drift, rather than from relatively recent founder effects.

Link

Y chromosomes and mtDNA of Sweden

It's interesting to see a study of the current population of a country. If one is interested in deep pre-historical origins, one must ensure that his sample has no known foreign ancestry among the lines of interest. But, if one is interested in the current and future course of the population, then the totality of the population must be sampled.

This is a good opportunity to track shifting gene frequencies due to immigration and/or differential fertility. It would be a good idea for countries to start spending some money on a genetic census of their population. This would not need to involve all the inhabitants, and could be carried out for the fraction of cost that governments pay to collect all sorts of other statistics. Such a census would provide an important source of data to future scientists investigating the demography of Europe during this transitional era.

From the paper:

Several haplogroups had interesting frequency patterns, but also wide confidence intervals, necessitating caution in the interpretations. The mtDNA haplogroup with the strongest geographical cline, U5b, is known to have high frequencies among the northern Saami population, consistent with our results (Tambets et al. 2004). The high frequency of the Y-chromosomal R1b in the south, observed also by Karlsson et al. 2006; is consistent with its abundance in Central Europe and Denmark (Semino et al. 2000; Brion et al. 2005). Haplogroup R1a1 is more common in Norway than in Sweden (Dupuy et al. 2006), and its high frequency in Värmland/Dalarna and Halland supports the historically plausible connection to Norway (Lindqvist, 2006). Y-chromosomal haplogroup N3 (Lappalainen et al. 2006) and mtDNA haplogroup H1f (Loogväli et al. 2004; Lappalainen et al. 2008) are common in Finland, and had increased frequencies in several Swedish counties with historical ties to Finland: Eastern Svealand was the most important destination of the Finnish immigration wave in the 1970's; in Norrland the Finnish influences date back to ancient times and in Dalarna to the 17th century (Pitkänen 1994).

When compared to previous knowledge of ethnic Swedes without immigration in their familial background (Lappalainen et al. 2008), the frequencies of several haplogroups showed effects of 20th century immigration from more distant countries. The Y-chromosomal I1a had decreased frequencies in Malmö and Gothenburg most probably due to replacement by haplogroups that are common among immigrants. African immigration contributes to the frequencies of mtDNA haplogroups L3*(xN,M) and L* (xL3) (Chen et al. 2000), and Y-chromosomal haplogroup A (Underhill et al. 2001; Jobling & Tyler-Smith 2003), while Near Eastern influence can be seen in mtDNA haplogroup U7 and possibly J (Richards et al. 2000; Abu-Amero et al. 2007; Achilli et al. 2007). Asian and American immigration can be observed in the slightly elevated frequencies of mtDNA haplogroups M, A, C, D and G (Quintana-Murci et al. 2004; Hill et al. 2007) and the Y-chromosomal O, K* and P* (Underhill et al. 2001; Jobling & Tyler-Smith 2003). The frequency of the Y-chromosomal haplogroup I1b may associate to immigrants from Balkan and Eastern Europe (Rootsi et al. 2004). In Malmö and Gothenburg immigration was the main contributor to their isolated positions in the Y-chromosomal PCA plot and probably also to the higher diversities compared to the surrounding populations. These phenomena were not observed in Stockholm, where most of the immigrants come from Finland (Statistics Sweden, http://www.scb.se).

Annals of Human Genetics doi: 10.1111/j.1469-1809.2008.00487.x

Population Structure in Contemporary Sweden—A Y-Chromosomal and Mitochondrial DNA Analysis

T. Lappalainen et al.

Abstract

A population sample representing the current Swedish population was analysed for maternally and paternally inherited markers with the aim of characterizing genetic variation and population structure. The sample set of 820 females and 883 males were extracted and amplified from Guthrie cards of all the children born in Sweden during one week in 2003. 14 Y-chromosomal and 34 mitochondrial DNA SNPs were genotyped. The haplogroup frequencies of the counties closest to Finland, Norway, Denmark and the Saami region in the north exhibited similarities to the neighbouring populations, resulting from the formation of the Swedish nation during the past millennium. Moreover, the recent immigration waves of the 20th century are visible in haplogroup frequencies, and have led to increased diversity and divergence of the major cities. Signs of genetic drift can be detected in several counties in northern as well as in southern Sweden. With the exception of the most drifted subpopulations, the population structure in Sweden appears mostly clinal. In conclusion, our study yielded valuable information of the structure of the Swedish population, and demonstrated the usefulness of biobanks as a source of population genetic research. Our sampling strategy, nonselective on the current population rather than stratified according to ancestry, is informative for capturing the contemporary variation in the increasingly panmictic populations of the world.

Link

Vikings abroad in search of women

Viking Age Triggered by Shortage of Wives?:

Sept. 17, 2008 -- During the Viking Age from the late eighth to the mid-eleventh centuries, Scandinavians tore across Europe attacking, robbing and terrorizing locals. According to a new study, the young warriors were driven to seek their fortunes to better their chances of finding wives.

The odd twist to the story, said researcher James Barrett, is that it was the selective killing of female newborns that led to a shortage of Scandinavian women in the first place, resulting later in intense competition over eligible women.

...

Soren Sindbaek, assistant professor of medieval and Renaissance archaeology at Denmark's University of Aarhus, told Discovery News that the new paper "is very right in pointing out the inadequacy" of former explanations for the Viking Age.

"We need indeed to seek for an individual, social motivation behind the fact that a large number of young men chose to set out on extremely risky voyages in hopes of acquiring wealth and esteem in foreign lands," Sindbaek said.

ISBA3 abstracts

Many abstracts from the International Symposium on Biomolecular Archaeology.

Various DNA / Technology

Human ancient DNA analysis within The Genographic Project: a project update and preliminary results from two powerful multiplex SBE typing methods

Wolfgang Haak1, Juan J Sanchez2, Clio Der Sarkissian1, Christina Adler1 & Alan Cooper1

1 The Australian Centre of Ancient DNA, School of Earth & Environmental Sciences, The University of Adelaide, North Terrace Campus, SA-5005 Adelaide, Australia
2 National Institute of Toxicology and Forensic Science, Canary Islands Delegation, 38320 Tenerife, Spain.

The Australian Centre of Ancient DNA (ACAD) is one of 11 Regional Centres of The Genographic Project (TGP), and the only centre dedicated solely to ancient human DNA analyses. Our aim is to provide a temporal perspective to the movements and presence of prehistoric and historic populations through ancient DNA, and establish a time depth to the detailed genetic landscape being generated from the largescale modern human population data of TGP. We will present an update on the current activities and an overview of the protocols and strategies used in the ACAD. The retrieval of authentic human ancient DNA is plagued by methodological problems, and to deal with these we are relying on state-of-the-art methods ranging from sample collection through to data generation. We are using SPEX and multiplex PCR assays followed by SBE typing to analyse both mtDNA coding region markers and nuclear NRY markers. New multiplex assays were designed to amplify highly degraded DNA with an average amplicon length of 60-80bp, targeting 22 mtDNA SNPs and 25 NRY SNPs - to match the core marker panel used within TGP. Preliminary results show that the SBE typing protocols are robust and prove to be highly efficient in targeting minute amounts of suriving aDNA. In addition, the main advantage of SBE has proven to be the detecting power of omnipresent (background) contamination. We conclude that the mtDNA and NRY SBE assays, in combination with sequence data from the mtDNA control region (backed up by cloning and SPEX), and specialised sample collecting systems, provide a powerful means to effectively generate largescale (pre-)historic population data from ancient human samples.

What colour was Attila the Hun’s horse?: genetic signatures of phenotypic traits in archaeological materials

Mim A. Bower¹, Michael G. Campana², Diane Lister¹, Mark Whitten³, Kathy M. Dominy⁴, Angela M. Murphy⁵, Paula Jenkins⁶, Richard Sabin⁶, Michael Akam⁷, Robert Asher⁷ & Matthew Binns⁵.

¹ McDonald Institute for Archaeological Research, University of Cambridge, Downing Street, Cambridge, CB2 3ER, UK.

² Department of Archaeology, University of Cambridge, Downing Street, Cambridge, CB2 3DZ, UK.

³ Comparative Population Linguistics Group, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany.

⁴Department of Medical and Molecular Genetics, King's College London School of Medicine, Guy's Hospital, London, SE1 9RT, UK.

⁵Department of Veterinary Basic Sciences, the Royal Veterinary College, Royal College Street, London, NW1 0TU, UK.

⁶ Department of Zoology, Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD, UK

⁷Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.

Studying the phenotype, i.e. what a plant or animal looked or behaved like, is perhaps the next “Holy Grail” in ancient DNA research. If we could trace the genes responsible for particular characters in archaeological specimens, it would open the door to some highly relevant and interesting questions in archaeology; the least of which may be the elusive “domestication gene”, but could cover more basic questions, such as: did people in mediaeval Europe prefer cows that had a tendency to lay down fat or muscle? When and where did a particular genetic disorder enter a domestic population? Was the spread of agriculture influenced by adverse environmental conditions? Or what colour was Attila the Hun’s horse? We could begin to understand past approaches to animal and plant husbandry, the expression of past human choice and selection, and past understanding of biological processes such as heredity. Fortunately, molecular biology is fast uncovering the genes responsible for particular phenotypic traits – the only problem, and a significant one perhaps, is looking for these genetic signatures in ancient DNA, known to be a recalcitrant material for analysis.

Here we present the results of our research on phenotypic loci in various historic and archaeological materials (bones, teeth, parchment, hide, seeds), and discuss the potential for future of phenotypic research in archaeogenetics.

Greece & Italy

Aristophanes and Stable Isotopes: Comparing literary and isotopic evidence of diet in Classical Thebes, Greece.

Efrossini Vika ^1,2, Mike Richards^3,4, Holger Schutkowski² and Vassilis Aravantinos⁵

¹ School of Conservation Sciences, Bournemouth University, BH12 5BB, UK

² Division of Archaeological, Geographical and Environmental Sciences, University of Bradford, BD7 1DP, UK

³ Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, DE

⁴ Department of Archaeology, Durham University, DH1 3LE, UK

⁵ IX Ephorate of Prehistoric and Classical Antiquities, Thebes 60200, GR

The greatest advantage for an archaeologist working with historically dated material is the existence of literary sources, which can complement scientific analyses in archaeology. The present study compiles a dietary profile for the inhabitants of Classical Thebes, using δ¹³C and δ¹⁵N isotope analysis, comparing this evidence with information on diet as presented in Aristophanes’ comedies (Acharnians, Plutus, Wasps).

Aristophanes’ work is abundant in satirical scenes, which emphasize personal attributes and local characteristics. Within this realm, it is possible to extract information on dietary habits, trade and economy in the Classical times. Merchants from Thebes oftentimes appear in his work, reflecting the city’s wealth. Among the goods deriving from this region, eels feature prominently, and are praised as an exceptional delicacy.

Stable isotope analyses of bone collagen were carried out for individuals from the Classical burials of the Northeastern cemetery of Thebes. Results show a remarkable increase of the nitrogen values relative to the previous periods, which is not accompanied by an analogous enrichment of the carbon values. This profile can partly be explained by an increased consumption of freshwater sources in Thebes during the Classical times, such as Aristophanes’ famous eels.

The results demonstrate how the integration of isotopic and literary evidence can provide novel information about Classical society in Thebes.

Greek myths

Terry Brown
Faculty of Life Sciences, Manchester Interdisciplinary Biocentre, University of Manchester, M1 7DN, UK

The Greek Bronze Age is the time of the Homeric legends and the Greek myths. The work of Schliemann, Tsountas, Wace and others revealed the material remains of the Mycenaean civilisation that was dominant in the Aegean during the 17th to 12th centuries BC, these remains including human skeletons, some buried with rich grave goods. Biomolecular archaeologists have been attracted to the Greek Bronze Age because there are interesting kinship questions for the burials at several sites, and because aDNA could throw light on the impact of diseases such as malaria on these societies. Addressing these questions clearly requires that aDNA is preserved in the relevant material, and our conclusion after ten years of work is that largely it is not. We have found no indication of aDNA at Lerna, Antron Grave Circles A and B and Mycenae Grave Circle A. At Mycenae Grave Circle B, we detected mitochondrial aDNA in just four of the 22 skeletons that we studied. Only at Kouphovouno have we have obtained sufficient aDNA results to attempt any kind of archaeologically relevant study. The distinguishing feature of Kouphovouno is that we obtained skeletal samples immediately after their excavation. We used optimised PCR systems in order to maximise our chances of detecting aDNA if it was present, but we also used a high containment facility and took scrupulous care to remove surface contamination from the bone samples and to prevent cross-contamination with PCR products from previous experiments. We also confirmed that our negative results were not due to inhibition of PCRs by substances co-purifying with aDNA. Negative results tend not to get widely publicised – we would have preferred positive ones but not if they lead to new Greek myths.

Late Bronze Age Diet in the Greek Peloponnese

E.I.Petroutsa1 & M.P.Richards2

1.20 Koundouriotou str., Exarcheia, 10683 Athens, Greece

2.Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany


In this paper we present the results of an isotopic study of bone collagen samples from four Late Bronze Age cemeteries from the Northern Peloponnese in Greece: Voudeni, Aghia Triada, Almyri and Kritika. Through isotope analyses we sought to characterise the general diets in these four sites, especially the amounts of marine protein, as well as animal vs. plant proteins in diets. We also compared the isotopic results from these sites with other Bronze Age sites, including Mycenae. Despite the coastal location of most of the sites we could not find evidence of any significant consumption of marine foods. Instead, most human diets are based on a mixture of plant and animal protein, from C3 terrestrial resources.

Preliminary results of C and N isotope analyses and 14C dating of prehistoric humans and animals from the Mesolithic-Neolithic site of Grotta dell’Uzzo, Sicily, Italy

Marcello A. Mannino1, Sahra Talamo1, Rosaria Di Salvo2, Vittoria Schimmenti2, Marcello Piperno3, Sebastiano Tusa4, Antonio Tagliacozzo5, Michael P. Richards1,6

(1) Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig (Germany)

(2) Museo Archeologico Regionale ‘Antonino Salinas’, Via Bara all’Olivella 24, 90133 Palermo (Italy)

(3) Dipartimento di Scienze Storiche, Archeologiche ed Antropologiche dell’Antichità, Sezione di Paletnologia, Università di Roma ‘La Sapienza’, Via Palestro 63, 00185 Roma (Italy)

(4) Soprintendenza del Mare, Palazzetto Mirto, Via Lungarini 9, 90133 Palermo (Italy)

(5) Museo Nazionale Preistorico Etnografico ‘L. Pigorini’, Piazzale Guglielmo Marconi 14, 00144 Roma E.U.R. (Italy)

(6) Department of Archaeology, University of Durham, South Road, DH1 3LE Durham (United Kingdom)

Grotta dell’Uzzo is one of the key sites in the Mediterranean for the study of the changes in subsistence that took place in the transition from hunter-gatherer (Mesolithic) to agro-pastoral (Neolithic) economies. The cave is also important because 13 Mesolithic humans have been unearthed from 11 burials excavated within it. In order to study the diets of these humans, C and N isotope analyses were undertaken on skeletal remains recovered in the burials and from different trenches excavated at the cave. The preliminary results of these analyses suggest that the main sources of dietary protein were terrestrial and probably originated from the consumption of mammalian herbivores such as red deer (the most commonly exploited animal for much of the cave’s occupation). The contribution of marine resources to human diet at Grotta dell’Uzzo was probably not very significant in absolute terms. This finding might appear to be in contrast with the results of the archaeozoological studies, which have demonstrated that there was an intensification in shellfish collecting and in fishing from the end of the Mesolithic to the inception of the Neolithic. However, given that on the basis of the chronological data currently available the burials predate the end of the Mesolithic, the results of the isotope analyses are in line with the findings of the archaeozoological studies and of the seasonality studies, which have shown that marine resources (mainly represented by shellfish) were not exploited throughout the year before the final stages of the Mesolithic.

Plants

Phylogeographic analysis of barley (Hordeum vulgare) landraces shows that the distribution of lineages retains an imprint from the initial patterns of agricultural spread through Europe.

Huw Jones1, James Cockram1, Lydia M Smith1, Ian MacKay1, Robin G Allaby2, Terrence A Brown3, Wayne Powell1

1 National Institute of Agricultural Botany, Huntingdon Road, Cambridge, CB3 0LE

2 Warwick HRI , Wellesbourne, Warwick , CV35 9EF

3 Faculty of Life Sciences, Manchester Interdisciplinary Biocentre, 131 Princess Street, Manchester, M1 7DN

In the 8000 years since barley was first introduced into the continent of Europe, the evolution of this crop species has been subject to selection under biotic and abiotic pressures. European farmers will have influenced the distribution of barley ecotypes by the movements of early farmers and by their selection of preferred types. The evolution of barley in Europe will also reflect the differing environments in which it has been grown. Modern barley cultivars from across Europe can be seen to differ in their morphology and growth habits. We have attempted to dissect the origins of these different forms by examining the landraces and assess the relative importance of human and environmental selection on modern barley lineages.

We have sampled barley landraces from Europe and discovered their population structure by microsatellite genotyping and statistical analysis. We have used passport data from germplasm collections to characterise the 2-row / 6-row head morphology, hulled / naked grain morphology and the spring / winter growth habit of these sub-populations. The genetic variation underlying key adaptive traits controlling flowering time has been explored by re-sequencing the photoperiod response gene Ppd-H1 and by haplotype analysis at the spring / winter vernalisation genes Vrn-H1 and Vrn-H2. These studies were designed to run alongside analysis of ancient DNA and historic DNA from barley and emmer wheat collected from across Europe.
The population structure we have discovered divides barley landraces into a number of sub-populations each with a distinct geographic distribution. Our genetic data for key adaptive traits allows us to understand the environmental influence on the geographic distribution of each lineage. Where lineages with a similar adaptive profile have distinct geographic distributions we see the imprint of early dispersal by ancient farmers.

Stable isotope evidence for the consumption of millet in Bronze Age Italy

Mary Anne Tafuri1, Oliver Craig2 & Alessandro Canci2
1 Dipartimento di Biologia Animale e dell’Uomo, Sapienza Università di Roma, P.le A. Moro, 5, 00185 Roma. Italy – email: maryanne.tafuri@uniroma1.it

2 Department of Archaeology, University of York, BioArch Biology, S Block PO Box 373 York YO10 5YW, UK

3 Dipartimento di Storia e Tutela dei Beni Culturali, Università degli Studi di Udine, Via Palladio, 8, 33100 Udine. Italy


This study presents, via carbon and nitrogen stable isotope analysis on human and animal bone collagen, new data on diet and subsistence strategies at northern and southern Italy Early and Middle Bronze Age sites, which clearly indicate the direct or indirect consumption of C4 plants. On the basis of paleobotanic data available and as suggested by previous similar studies, we argue here that the isotopic signal obtained can be associated with the consumption of millet (P. miliaceum and Setaria italica). If such an interpretation were true, while we wait for further paleobotanical and isotopic studies, we should consider the results obtained as the earliest evidence of millet consumption in prehistoric Europe. We thus suggest a possible pattern of distribution in the Peninsula of the practice of production and consumption of millet, while setting a new agenda on food security and subsistence strategies in prehistoric Italy

Cattle & Goats

A PCR system free of contaminating DNA for the amplification of bovine DNA from bovine fossils

Camille Berthelot, Sophie Champlot, Marie Liouville, Thierry Grange, Eva-Maria Geigl

Institut Jacques Monod CNRS UMR 7592, Universités Paris 6 et 7, Tour 43, 2, Place Jussieu, 75251 Paris cedex 05, France

Palaeogenetic analyses of bovine bone remains from many Neolithic sites in Europe and in Southwest Asia suffer from poor DNA preservation in these bones that increases the risk of amplification of contaminating modern bovine DNA. Indeed, trace amounts of contaminating bovine DNA occur ubiquitously. In particular, they can be found at low quantities in biochemical reagents used to extract and amplify DNA. These contaminating molecules mimic ancient DNA molecules. Indeed, the contamination rate often resembles the success rate of ancient DNA studies from bovine remains and the length of the contaminating DNA fragments is often comparable to ancient DNA fragments. We elaborated a decontamination protocol for PCR reagents combining various treatments to reduce contamination towards zero. This system significantly increases the reliability of ancient DNA results from bone remains of domesticated animals.

Detecting selection in ancient cattle remains: Pre industrial selection in Bos Taurus and SNP typing in medieval cattle remains

Emma Svensson1, Anders Götherström1

1 Evolutionary Biology, Evolution Genomics & Systematics, Uppsala University, 752 36 Uppsala, Sweden

Historic and prehistoric animal breeding is an enigmatic topic, complicated to approach with conventional genetics and osteology. Questions like when it started, and how strict it was, are of general interest, but it is also complicated to generate a suitable dataset for such questions. By tracing changes in genetic diversity with serial data we can find out how cattle has changed since the domestication of the aurochs to become the array of breeds seen today. Cattle are likely to have been subjected to selection predating the 18th century but the information is scarce. Using a 12plex SNP stream system alongside pyrosequencing we typed up to eight coding and six neutral SNPs in 142 ancient and 216 modern Bos Taurus from Northern Europe. We found a significant decrease in total heterozygosity over time for the coding SNPs which are presumably associated with phenotypic traits such as milk quality and coat colour while neutral markers on the other hand don’t show any significant change over time. This suggests that the decline in diversity is caused by artificial selection and not other genetic processes. The medieval period was a dynamic time in northern European history. The society was moving toward a higher degree of specialization in general, and a number of towns based on trade arouse in Scandinavia. Our findings of early selection fit well with the more sophisticated farming and higher degree of animal breeding that likely occurred at this time.

Using new and old approaches to study bovid systematics and evolution across Eurasia

Alan Cooper¹, Kefei Chen¹, Beth Shapiro²

¹ The Australian Centre of Ancient DNA, School of Earth & Environmental Sciences, The University of Adelaide, North Terrace Campus, SA-5005 Adelaide, Australia

² Department of Biology, The Pennsylvania State University, 326 Mueller Laboratory, University Park PA 16802, USA

Ancient DNA studies of bovid remains from Europe have detected four main taxa: Bison bonasus (the European Bison); Bison priscus (Steppe bison), Bos primigenius (Aurochs); and early Bos taurus (Daisy). Studies of bones recovered from caves in the Urals and Caucasus, and from material dredged from the North Sea, have revealed a fifth European bovid – the Caucasus bison. Previously recognised only as a sub-species of European bison, this taxon appears to represent a separate species, with more genetic diversity than Beringian populations of Bison priscus, suggesting a long evolutionary history and stable population size. It has changed ecological dominance with Bison priscus at several points in the Pleistocene, which appear to be related to climatic and environmental change.

We have been using emulsion PCR and high-throughput hybridisation-based SNP screening systems that can simultaneously analyse 50,000 bovid SNPs to explore the genomic evolution of ancient bovids during the Pleistocene and subsequent domestication. We have been concentrating on pre-domestic Bos taurus specimens, as well as representatives of the other Pleistocene bovid species. This approach holds enormous promise for fine-scale temporal analyses of evolution in response to climate and environmental change, as well as archaeology and domestication.

The process of cattle domestication during the Neolithic as revealed by a large-scale palaeogenetic study

Eva-Maria Geigl, Mélanie Pruvost, Marie Liouville, Camille Berthelot, Reinhard Schwarz, Sophie Champlot, Thierry Grange, Virginia Bessa-Correia, Hans-Peter Uerpmann, Lamys Hachem, Hitomi Hongo, Séverine Braguier

Institut Jacques Monod CNRS UMR 7592, Universités Paris 6 et 7, Tour 43, 2, Place Jussieu, 75251 Paris cedex 05, France

Several disciplines can contribute to the elucidation of the processes of animal domestication during the Neolithic, such as archaeology, archaeozoology, and, more recently, isotope and genetic studies. The processes of domestication leave genetic signatures in the genomes of the domesticated animals that can be explored via the combination of both genetic analyses of extant domesticates and palaeogenetic analyses of bone remains of the first generations of domesticated animals and of their wild ancestors. We adopted this approach to shed light on the domestication of the aurochs. We studied roughly 250 Bos bone remains from Southwest Asia, according to archaeological and archaeozoological evidence the presumed centre of cattle domestication, and from France, the region where the two Neolithic migration currents mingled. To obtain authentic palaeogenetic results, several methodological difficulties related to poor DNA preservation and reagent contamination had to be solved. We will present both the methodological challenge that we encountered and overcame and the results of our large-scale study.

Cattle domestication and the troublesome aurochs

Cecilia Anderung1, Jurgita Baubliene2, Daniel Makowiecki3, José Miguel Carratero4, Linas Daugnora2, Juan Luis Arsuaga5 and Anders Götherström6

1Palaeontology Department, Natural History Museum, Cromwell road, London SW7 5BD, Great Britain.
2Department of Anatomy and Histology, Lithuanian Veterinary Academy, Tilžes str. 18, LT-3022, Kaunas, Lithuania
3Institute of Archaeology, Nicolaus Copernicus University, Podmurna 9/11 87-100 Toruń, Poland
4Laboratorio de Evolución Humana, Departamento Ciencias Históricas y Geografía Edificio I+D+I Plaza de Misael Bañuelos s/n, 09001, Burgos, Spain
5Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos c/ Sinesio Delgado Nº 4 Pabellón 14, 28029 Madrid, Spain
6Department of Evolutionary Biology, Evolutionary Biology Centre, Uppsala University, 75236 Uppsala, Sweden

A number of genetic studies relating to the origin of modern cattle have been published recently. In particular the extinct European aurochs (Bos primigenius), progenitor of the modern domesticated form, has attracted a lot of attention. European aurochs bones that have yielded mitochondrial DNA are genetically clearly different from modern cattle. This genetic data, in combination with the genetic patterns observed in modern cattle, has led to the suggestion that modern cattle have a single origin in the Near East, and that the European aurochs population had little to do with their domestication.

However, we ask the question: could this picture of European cattle domestication be too simple?

Here we present results from an investigation of published and novel aurochs sequences from Lithuania, Spain, Poland, and Britain, giving special attention to the Spanish sequences. We looked at the mitochondrial DNA variation in Iberian aurochs remains and searched for aurochs sequences in a domestic context. We find that cattle and aurochs mitochondrial sequences from Iberia deviate from the pattern observed in sequences from Central and Eastern Europe.

In the light of this data, we discuss the possibility of local aurochs domestication events in Europe.

High mtDNA diversity among cattle and goats from the earliest Neolithic settlements on the European continent

Amelie Scheu1,2, Norbert Benecke2 and Joachim Burger1
1 AG Palaeogenetik, Institut für Anthropologie, Johannes Gutenberg-Universität, 55099 Mainz, Germany

2 Deutsches Archäologisches Institut, Eurasienabteilung, 14195 Berlin, Germany

The process of domestication includes a decline in genetic variability. Additional homogenisation occurs due to subsequent colonisation events, such as the Neolithisation of Europe. Our previous studies have shown genetic uniformity even among early Neolithic European cattle (Bollongino et al. 2006). But modern goats also share more than 90% of the same mtDNA haplogroup.

To find out exactly when and where this genetic bottleneck arose during the Neolithisation of Central Europe, we investigated remains of early domesticates on the border between Asia and Europe, i.e. at the origin of the trans-Danubian route of Neolithisation. That region, particularly the area around the Bosphorus and the transit country of Bulgaria, plays a crucial role.

We found higher mtDNA diversity among Neolithic and Bronze Age domesticated cattle East and West of the Black Sea (haplogroups T, T2 and T3) than in Central European populations. Among goats, we found the two different mtDNA haplogroups A and G. G had previously been found among modern goats only near the Fertile Crescent (Naderi et al. 2007).

Our results argue for large and genetically more diverse herds imported to this area and/or for intense trade. Furthermore, they indicate that a second wave of expansion in the direction of Central Europe is responsible for the final loss of mtDNA diversity.

Ice man Schnidi’s trousers: insight into prehistoric goat diversity

Angela Schlumbaum1, Serge Volken 2, Marquita Volken 3, Jörg Schibler4, Peter Suter 5 Kathrin Glauser6 & Albert Hafner 7

1 Institute of Prehistory and Archaeological Science, University of Basel, Spalenring 145, 4055 Basel, Switzerland

2 Gentle Craft, Rue du Rôtillion, 10, 1001 Lausanne, Switzerland

3 Gentle Craft, Rue du Rôtillion, 10, 1001 Lausanne, Switzerland

4 Institute of Prehistory and Archaeological Science, University of Basel, Spalenring 145, 4055 Basel, Switzerland

5 Archaeological Service of the Canton Bern, Brünnenstrasse 66, 3001 Bern, Switzerland

6 Archaeological Service of the Canton Bern, Brünnenstrasse 66, 3001 Bern, Switzerland

7 Archaeological Service of the Canton Bern, Brünnenstrasse 66, 3001 Bern, Switzerland
Since 2003 more than 300 prehistoric remains were discovered in the vicinity of a melting ice patch of the Schnidejoch (2750 m; Bernese Alps, Switzerland), paralleling the finds accompanying the Iceman from the Tisenjoch (Oetztal Alps, Italy/Austria; “Oetzi”). One leg of a Neolithic leather trouser was found and 14C dated to 2900 – 2600 BC. The morphological identification of the animal skin was ambiguous because of the bad preservation of diagnostic features. Based on grain patterns of the skin the leather was made either from sheep or goat.

Because of the importance of the object, a genetic identification of the animal species was attempted. After DNA extraction with QiAmp DNA Mini Kit a 70bp fragment of the mitochondrial cytochrome b gene was amplified in the diluted extract. The leather was made of goat skin (Capra hircus). Six geographically broadly distributed goat lineages are recognized based on mitochondrial d-loop variation, of these lineage A and C were identified by others in prehistoric goat bones from France. The “trouser’s goat” however, belongs to lineage B, which is common in Asia, but extremely rare in Europe today.

An Ancient DNA study from The Farm Beneath the Sand

Martin Bay Hebsgaard1

1 Dept. of Biology, University of Copenhagen, Universitetsparken 15, Denmark
Applying ancient DNA techniques on samples from the archaeological site “The Farm Beneath the Sand” (GUS) near Nuuk in Southwest Greenland is the first attempt to extract DNA from these relative young but novel samples. The sample site highlight the dramatic landscape changes that resulted in floodplain aggradations that eventual buried the site fixing the site under perma-frozen conditions.
The Farm beneath the Sand is situated on a plain surrounded by low mountains ca. 80 km east of Nuuk. When the building remains were found they were overlain by ca. 1,5 m thick layers of sand and gravel, and today the plain in front of the farmhouse appears as a sandy dessert intersected by meandering watercourses that are draining off the icecap.

All together the samples yielded DNA from humans, cattle, sheep, goat and reindeer. Quantification shows approximately 16 times more DNA from cattle than from sheep. Goat DNA was undetectable using Quantitative PCR. The amount of cattle DNA declines over time while sheep DNA probably reflects background variation.
Thanks to thick layers of sand and gravel that may have protected the DNA in the anthropogenic layers ancient DNA is usable in an archaeological context in the reconstruction of the past. In this example ancient DNA research has helped to refine and define archaeological interpretations of the Norse life by adding information not seen by the naked eye. In the future ancient DNA have the capacity to be used more vigorously to investigate the diet of the Norse and show what function specific farms may have had during the settlement. As in this study the future research is not limited to animal DNA but DNA from plant and humans can be used to address different questions.

Horses

Pleistocene Horses genetics before and after the last glacial maximum

Sebastian Lippold¹ and Michael Hofreiter¹

¹ Dept. Evolutionary Genetics, MPI for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig, Germany

We are investigating how the last glacial period affected the genetic composition of populations, particularly the horse (Equus spec.). Vast climatic changes that occurred between 30,000 and 12,000 years BP resulted in dynamic mammalian population structures. Restriction and expansion events during this period influenced both the genetic distribution and variability of a variety of mammals. We focus on western European horse populations, and try to characterize their genetic diversity and phylogeographic patterns both before and after the last glacial maximum. We sequence 600 bp of mtDNA from the mitochondrial D-Loop of different individuals obtained from different locations. Our initial results indicate tremendous genetic diversity, but no phylogeographic pattern within this marker. The genetic distribution of these ancient samples also falls within the broad diversity range apparent in recent horses. Because of this finding, we started screening the DNA samples for additional nuclear markers. Our comparison of these different markers has enabled us to reconstruct several scenarios for horse population dynamics during this period.

Investigating Eneolithic horse exploitation in northern Kazakhstan, via compound-specific stable carbon and deuterium isotope analysis of pottery.

Natalie A. Stear1, David Chivall1, Alan K. Outram2 and Richard P. Evershed1 1Organic Geochemistry Unit, School of Chemistry, University of Bristol, Clifton, BS8 1TS, UK 2 Department of Archaeology, SoGAER, University of Exeter,Laver Building, North Park Road, Exeter, EX4 4QE, UK The Eneolithic site of Botai in northern Kazakhstan has been the centre of much debate regarding its role in early horse domestication (Levine, 1999; Olsen, 2003). The faunal assemblage from this remarkable site was almost entirely comprised of horse (99%); however it has remained unknown whether any of these horses were domesticated. Horse domestication is extremely difficult to detect morphologically from skeletal remains and consequently it is necessary to establish a reliable proxy for detecting ancient domestic horse populations. Fermented mare’s milk (Kumyss) is commonly consumed in rural communities in Kazakhstan, a tradition dating back to prehistory. It is not clear if the milking of horses began (i) in the Eneolithic, during the height of the horse centred communities of the Botai culture; (ii) in the Bronze Age, in response to the milking of ruminants or, (iii) much later. If mare’s milk were identified in ancient pottery it would serve as conclusive evidence for the presence of domestic horses and enable a chronology of horse exploitation to be established. Equine fat residues can be identified in potsherds using compound-specific stable carbon isotope analysis, but unlike ruminant fats, equine milk and adipose fats are indistinguishable from one another based on δ13C values. However, we show that it is possible to further classify equine fats as either milk or adipose, based upon the δD values of their C16:0 and C18:0 fatty acids which are determined using GC-thermal conversion-IRMS (GC-TC-IRMS). This new proxy has been applied to organic residues extracted from potsherds from Botai as part of a large scale investigation of Eneolithic and Bronze Age pottery from sites in northern Kazakhstan. Using the δ13C and δD values obtained from the Botai residues we have been able to detect equine milk residues preserved within the pottery and consequently, we provide the first direct evidence for the presence of domestic horses at Botai during the Eneolithic.

Anatolia

Detecting dairying with stable calcium isotope ratios (δ44/42Ca) of bones and teeth

Linda M. Reynard^1,2, Robert E.M. Hedges¹ & Gideon M. Henderson²

¹ Research Laboratory for Archaeology and the History of Art, University of Oxford, Dyson Perrins Building, South Parks Road, Oxford, OX1 3QY, United Kingdom

² Department of Earth Sciences, University of Oxford, Parks Road, Oxford, OX1 3PR, United Kingdom

The use of dairy products in antiquity is an important part of the development of agriculture and pastoralism in Eurasia. They offer advantages of more efficient land use, improved nutrition, and more reliable and constant access to protein. Understanding the adoption of dairy, its timing, and geographical spread is relevant to developing a fuller understanding of changes in subsistence from the Neolithic.

Detecting and quantifying dairy product consumption in antiquity has, to date, relied on indirect evidence such as the age and sex structure of faunal remains and potsherd lipid residues.. To complement these methods, we have measured stable calcium isotope ratios (δ^44/42Ca) of bones and teeth which allows the direct detection of dairy consumption by prehistoric humans. Dairy products have lower δ^44/42Ca than other dietary calcium inputs, and this results in lower δ^44/42Ca of the dairy consumer. We have measured the δ^44/42Ca of human and animal bones from a range of archaeological sites by MC-ICP-MS. Results from the Mesolithic to the Neolithic at the key Near Eastern site of Abu Hureyra, Syria (11,100 –7,300 BP) show a δ^44/42Ca signal attributable to dairy consumption by ancient humans, with a changing pattern through time. Work on intra- and inter-tooth δ^44/42Ca variability is in progress as this material is expected to form a robust archive of in vivo isotope ratios.


Britain

An investigation into origins of individuals from a mass grave in Roman Gloucester, UK: strontium and stable isotope evidence

Carolyn Chenery^1,2, Gundula Müldner¹, Jane Evans², Louise Loe³, Nicholas Márquez Grant³, Hella Eckardt¹ Stephanie Leach¹, Mary Lewis¹

¹ Department of Archaeology, University of Reading, Reading, Berkshire.

² NERC Isotope Geoscience Laboratory, British Geological Survey, Keyworth, Nottingham.

³ Oxford Archaeological Unit Ltd, Janus House, Osney Mead, Oxford.

Contrary to popular assumptions, Britain under Rome was truly multi-cultural, with historical and epigraphic evidence recording the voluntary and forced migration of Gaulish, Germanic and North African individuals into the British provinces refs. This paper presents the results an isotopic investigation of population diversity in 1^st to the 4^th century Roman Gloucester; focusing on individuals found in a late 2^nd century mass burial pit and comparing them to those found in single graves.

The results suggest that the majority of the individuals buried in the London Road Cemetery were from areas within the UK. However, the isotope data has identified a number of individuals whose origins lay in a region with a warmer climate than the UK. Whether these were soldiers, their followers or merchants cannot be determined.

On Rome’s Northern Frontier: Multi-isotopic investigations into cultural diversity in Roman York

Gundula Müldner1, Carolyn Chenery1,2, Stephany Leach1, Mary Lewis1 & Hella Eckardt1

1 Department of Archaeology, University of Reading, Whiteknights, PO Box 227, Reading RG6 6AB, England

2 NERC Isotope Geoscience Laboratory, British Geological Survey, Keyworth, Nottingham, NG12 5GG, England

Britain under Rome was a multi-cultural society, with historical and epigraphic evidence attesting to the presence of migrants from continental Europe, North Africa and the Middle East. Here, we combine isotope analysis for the reconstruction of diet (C,N) and mobility (Sr, O) with skeletal (craniomorphometric) and archaeological data, in order to investigate diversity in Roman York, the political, military and administrative centre of the North. The results show a heterogenous population and highlight the varied life-histories of individuals in the northernmost provincial capital of the Empire; however, they also show that skeletal and isotopic evidence are not always easily correlated. It is suggested that a diet high in marine protein was used to demonstrate a “Roman” identity.

Investigating Marine Food Consumption in Prehistoric Humans via the δ13C values of Collagen Amino Acids

Philip Dunn1, Richard P. Evershed1 & C. Joshua Pollard2
1 School of Chemistry, University of Bristol, Organic Geochemistry Unit, School of Chemistry, Cantock's Close, Bristol, BS8 1TS, UK

2 Department of Archaeology and Anthropology, 43 Woodland Road, Clifton, Bristol BS8 1UU, UK

The current debate concerning the importance of marine foods in the diets of prehistoric peoples has stimulated interest in refining analytical approaches based on stable isotopes. An alternative to the widely bulk collagen approach is to investigate the stable carbon isotope composition of human bone collagen at the amino acid level. Our investigations of prehistoric humans from the Western Cape, South Africa, have shown that the Δ13CGlycine-Phenylalanine values for bone collagen amino acids correlate strongly with bulk δ15N values and provide a new proxy for marine food consumption. We are currently applying this new approach to human skeletal remains from northern Europe.
The tomb at Isbister lies on the south west coast of South Ronaldsay and is one of a number of Neolithic cairns found in the Orkney Islands. During excavation in 1976, the stalled main chamber and three side chambers were found to contain some 16,000 human bones and bone fragments deriving from a minimum of 341 individuals along with a wide range of faunal remains. Previous analyses of bone collagen from the human remains have shown that the tomb was in use from 4,500 to 3,800 BP and bulk collagen stable isotope values for 5 individuals lay in the range -19.9 to -21.2 ‰ indicating a predominantly terrestrial diet, which is surprising for people that lived so close to the coast.
We have now employed the recently introduced LC-IRMS technique to determine the δ13C values of bone collagen amino acids from 22 individuals from Isbister. The results will be compared to those obtained by GC/C/IRMS. The derived Δ13CGlycine-Phenylalanine values give enhanced insights into the dietary habits of the people of Isbister.

Northern Europe

Large-scale FLX-sequencing and the Swedish Neolithic

Helena Malmström1,2, Anna Linderholm3, M. Thomas P. Gilbert2, Mikael Brandström1, Jan Storå4, Petra Molnar4, Christian Bendixen5, Gunilla Holmlund6, Kerstin Lidén6, Anders Götherström1, Eske Willerslev2

1Department of Evolutionary Biology, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden

2Ancient DNA and Evolution Group, Biological Institute, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen O, Denmark

3Archeological Research Laboratory, Stockholm University, SE-106 91 Stockholm, Sweden

4Osteoarchaeological Research Laboratory, Stockholm University, SE-106 91 Stockholm, Sweden

5Department of Genetics and Biotechnology, Danish Institute of Agricultural Sciences Research Centre Foulum, K25 PO Box 50, DK-8830 Tjele, Denmark

6National Board of Forensic Medicine, Department of Forensic Genetics and Forensic Toxicology, Artillerigatan 12, SE-587 58 Linköping, Sweden

The relationship between the contemporary hunter-gatherer Pitted Ware Culture (PWC) and the farming Funnel Beaker Culture (TRB) in Middle Neolithic (3300-2500 B.C.) Sweden has been debated for more than a century. We approach this issue by determining the genetic signatures of skeletal remains from both complexes. Initially, we generated large amounts of “cloned” ancient mitochondrial DNA (mtDNA) PCR amplicons using a single run of the Genome Sequencher FLX System, and a recently described primer tagging protocol. Our data set consisted of Hypervariable Region I (HVRI) amplicons derived from bleach pre-treated powdered bone from Neolithic humans as well as from a large number of negative controls (animal samples, extraction and PCR blanks). We compared the ‘clone’ data with sample quality indicators, such as the number of PCR starting template molecules and the degradation ratio of DNA in the sample (number of long/short fragments). The data shows distinct patterns that differ between high and low quality extracts. After establishing the efficacy of the large scale sequencing approach, additional high-quality PWC and TRB samples (based on collagen preservation) were sequenced in a second FLX run. The compiled data yielded unambiguous HVRI sequences for approximately 40 Neolithic human samples, each compiled from cloned, duplicate PCR amplicons derived from overlapping HVRI fragments. F-statistics and AMOVA revealed significant genetic differences between the PWC and TRB samples, indicating that they indeed comprise of two distinct groups.

Allele frequencies of the lactase gene in Scandinavian Neolithic populations, hunter-gatherers vs. farmers

Anna Linderholm1, Helena Malmström2, 5, Love Dalén3, Kerstin Lidén1, Jan Storå4, Petra Molnar4, M. Thomas P. Gilbert5, Eske Willerslev5, Gunilla Holmlund6, Anders Götherström2
1Archeological Research Laboratory, Stockholm University, SE-106 91 Stockholm, Sweden

2Department of Evolutionary Biology, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden

3Marie Curie Fellow, School of Biological Sciences, University of London, United Kingdom

4Osteoarchaeological Research Laboratory, Stockholm University, SE-106 91 Stockholm, Sweden

5Ancient DNA and Evolution Group, Biological Institute, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen O, Denmark

6National Board of Forensic Medicine, Department of Forensic Genetics and Forensic Toxicology,Artillerigatan 12, SE-587 58 Linköping, Sweden

Genetics and culture are believed to interact, but it has been difficult to find direct evidence for the process. One example that has been put forward as a candidate is lactase persistance in adulthood, i.e. the ability to continue drinking milk. This genetic trait is believed to have evolved within a short space of time in connection with the emergence of farming cultures. Here we investigate certain Scandinavian Neolithic populations and their allele status with respect to the gene responsible for lactase persistance. We find that the allele responsible for lactase persistance was common and not significantly different from modern Swedish populations among Neolithic farmers, whereas Neolithic hunter-gatherers had a lower frequency of the allele.

aDNA analysis of human population samples from the Danish past – what have we learned?

L.C. Melchior¹, N. Lynnerup² and J.Dissing¹.

¹Research Laboratory, Institute of Forensic Medicine, University of Copenhagen, Denmark, ²Laboratory of Biological Anthropology, Institute of Forensic Medicine, University of Copenhagen, Denmark.

During an ongoing survey of the Danish genetic past we analyzed mtDNA from teeth from human remains from a range of burial sites from medieval times to the Stone Age. To ensure the highest possible degree of reliability generally accepted authentication criteria (including sequencing of multiple clones and replication by different researchers) as well as additional precautions (e.g. testing of laboratory performance) were observed.

Unequivocal assignment of mtDNA haplotypes was possible for more than 50 ancient subjects; however, the success rate varies substantially between sites. The highest success rate (11 out 11) was obtained with freshly excavated Viking Age subjects (ca AD 1,000), but good results were also obtained for recently excavated Iron Age sites (AD 0-400). Poor results were found with highly manipulated subjects (contamination) and with most of the Stone Age samples (4,500-5,000 YBP).

While laboratory related contamination can effectively be prevented by the abovementioned precautions, the most serious challenge to authenticity is caused by pre-lab contamination. This problem was tackled using several approaches including analysis of DNA damage patterns, haplotyping of archaeologists, phylogenetic testing and as the most efficient measure, sampling from the ancient subjects during exhumation. We show that reliable retrieval and analysis of DNA from ancient humans is indeed a possible undertaking.

A high degree of nucleotide diversity was observed in the ancient populations, and at four out of six locations the diversity was considerable higher than among modern Danes. Also, we observed a consistently higher abundance of Hg I (10-20%) than among modern Danes (~3%).The effect of the Black Death has been suggested as a possible explanation for a similar decline in the English genetic diversity. Interestingly, we found the highest genetic diversity in a 15^th century population sample long after the major outbreak in the14^th century.

NspI typed transition within PRNP gene (A385G / Met129Val) confirms rapid shift in allele frequencies during The Second Millennium

Henryk W. Witas1, Magdalena Kołodziejczak1, Paweł P. Liberski2
1 Dept. of Molecular Biology, Medical University of Lodz, 91-738 Lodz, Sporna 36/50, Poland

2 Dept. of Molecular Pathology and Neuropathology, Medical University of Lodz, 92-216 Łódź, Pomorska 251, Poland

Although precise biological role of prion proteins (PrPC) is still a subject of extensive study and debate, a few suggestions of their involvement in cellular processes have been described, including involvement in biology of synapse, short-term memory formation and long-term memory consolidation. Prion proteins became commonly recognizable as a cause of some human transmissible spongiform encephalopathies (TSE). Among a number of SNPs, A385G / Met129Val is assumed as a factor involved in the pathogenesis of TSEs (e.g. kuru) and a marker of memory efficiency as well. Although frequency of Met129Val alleles are precisely characterised for modern populations and its significance discussed, no data for historic and prehistoric populations to compare are available.

Specimens came from seven medieval cemeteries located throughout today’s Poland, and have been excavated recently. Teeth stored at low temperature underwent procedures generally accepted for aDNA isolation, performed automatically (MagNa Pure, Roche) at least two times on different teeth of each individual. Only the samples represented by collagen quantity above 2% dry weight, negative result of appropriate mock controls throughout isolation and amplification procedures, with successfully cloned (Amersham) and sequenced (AB 310) PCR products have been considered as authentic ancient templates. Moreover, we have applied NspI restriction analysis as a method for recognition and retrieval of undamaged ancient sequences.

The results show rapid increase in PRNP allele A frequency (Met 129) since the beginning of the Second Millennium (0.51 v. 0.65), accompanied by slight drop in heterozygotes (0.49 v. 0.39) and significant rise in Met homozygotes (0.27 v. 0.45).

As compared to present Polish as well as present European PRNP alleles frequency, medieval specimens provided the data which suggest altered mode of PRNP alleles transmission within last 35-40 generations. Although the nature of mechanism leading to observed changes is unclear, the impact of demographic factors is probably the most pronounced one affecting the process of local fluctuations of Met allele spreading out. However, the effect of selection processes should also be considered. This work is being supported by grant from Ministry of Science and Higher Education

Mice and Vikings

Eleanor Jones1
1 Department of Biology, University of York, Heslington, YO10 5DD, United Kingdom
Originally native to the northern Indian subcontinent and the Middle East, house mice (Mus musculus) have spread to their current near global distribution by exploiting a commensal niche with humans, originally in agrarian settlements. Mice also owe their current distribution pattern to human movements: they have reached the areas they now inhabit by being accidentally transported with grain and livestock foods. This close association between mice and humans means we can use information from the current genetic distribution of house mice to make inferences about past human colonisations and cultural linkages. In this study, we used mitochondrial DNA sequences from modern house mice in Great Britain, France, Ireland, Iceland, the Faeroe Islands and Norway to identify patterns in the their distribution, and tie these in to historic human migrations. The mice appear to be telling us about Norse Viking colonisations, and add a useful source of information to complement archaeological and historical data.

East Asia & Pacific

Genetic relationship of Human Skeletal Remains from an archaeological cemetery

Sang Hyun Jee 1, Yun Ji Kim 1, Yong Jae Chung1 & Min Seok Seo 1

1 Conservation Science Division, National Research Institute of Cultural Heritage, 472 Munji-dong, Yuseong-gu, Daejeon, 305-380, South Korea

We carried out genetic analyses of human skeletal remains from cemetery of a historic site, Myeong-arm-ri of Asan in South Korea. According to archaeological evidences, this site had been constructed from the Neolithic Age to the Joseon Dynasty. Twenty one human skeletons excavated from thirty pit tombs that have outer coffin build up into plaster dated to the Joseon Dynasty (14-19th century). To identify the genealogy and traditional burial pattern were assessed using mitochondrial DNA (mtDNA) and Y chromosomal STRs. We take cautious to avoid erroneous recombination by the segmental and modern contaminations were derived from researchers and all experimental stages. We sequenced the segmental amplicons of the hyper variable regions (HVRs) of mtDNA, and appointed relevant haplogroups according to the sequence polymorphism using the known mtDNA database. We also applied variable short tandem repeat (STR) marker in Y chromosome to understand paternal lineage and kinship among the burials. Especially, we interested in the four burying together and examined genetic relationship more closely between two individuals.

Not quite in the bag: A systematic bioarchaeological approach to the question of South American chickens origins

Greger Larson
1 Dept. of Archaeology, Durham University, South Road, DH1 3LE, UK

Though chickens were undoubtedly introduced into the American continents by the Spanish after their arrival in the 15th century, there is an ongoing debate as to the possible that Polynesians traveling across the Pacific introduced chickens to South America before Europeans did so. A recent publication concluded on the basis of ancient DNA extracted from an archaeological Chilean chicken bone that domestic fowl were present in a pre-Columbian context and that those chickens possessed a Polynesian genetic signature. In order to test this hypothesis, we generated mitochondrial DNA control region sequences from 41 modern, native Chilean specimens and analyzed them within a database consisting of both the published ancient DNA sequences and ~1,000 globally distributed modern domestic chicken sequences. Our modern Chilean sequences cluster closely with haplotypes predominantly distributed amongst European, Indian, and Southeast Asian chickens, consistent with a European genetic origin. The previously published, apparently pre-Columbian, Chilean specimen and seven pre-European Polynesian specimens, also cluster with the same European/Indian subcontinental/Southeast Asian sequences, providing, at this stage, no support for a Polynesian introduction of chickens to South America. Ancient DNA sequences from two archaeological sites on Easter Island, however, cluster with chickens found in Island Southeast Asia, and may represent a genetic signature of an early Polynesian dispersal as far as Easter Island. Lastly, we modeled the potential marine carbon contribution to the Chilean archaeological specimen (thus revising the derived date of the specimen) which cast doubt on the pre-Columbian age of the chicken remains. Definitive proof of a pre-Columbian introduction will require excavating more chicken bones, and further analyses of ancient DNA and radiocarbon data from Chilean and Polynesian archaeological excavations.

A New Bioarchaeological Clue for the DongHu Nationality

ZHANG Quan-chao, CHANG E，ZHU Hong

( Research Center f or Chinese Frontier A rchaeology , Jilin University , Changchun , Jilin , 130012 , China)

Abstract: DongHu nationality is a branch the Hu population who acted actively in the northeast of Yan. Whilst the remains of DongHu had not been confirmed for a long time, The tombs discovered in linxi Jinggouzi site in 2002 suggested a new clue for exploring the remains of Donghu in Chifeng area. These tombs not only meet the condionons in relation to DongHu in the aspects of time and region, but also conformed to DongHu characteristics of economic style and ethic features. In this article, human remains unearthed from the Spring and Autumn-Warring states cemetery at the Jinggouzi site in Linxi county, Inner Mongolia were studied. The morphological features of Jinggouzi group crania show that the racial type is closely related to the modern North Asiatic Mongoloids, and some physical characteristics of these skulls are closer to the ancient XianBei population in the north China and the modern Mongol. Ancient DNA sequences from ancient human remains have provided very important information on human evolution, blood relationship and migration, making ancient DNA research an important field of molecular anthropology. This study illustrates ancient DNA extraction, amplification and sequencing of five individuals of an ancient population buried in the west cemetery at Jinggouzi site in Inner Mongolia. A phylogenetic tree, a two-dimensional PC plot and MDS plot are constructed using mtDNA sequences from the ancient population and several modern Eurasian populations. However, the application of the techniques of ancient DNA allows us to explore the fasten source of the ancient population. Therefore, the genetic evidence raise the important meaning for the study of archeological culture in the east of Inner Mongolia during Spring and Autumn-Warring states. In addition, this new study which based on genetics and traditional archaeology on the development of populations from the north steppes of our country during Spring and Autumn-Warring states, provides precious data. In this study, we examine Jinggouzi population paleodiet using stable isotope ratios of carbon and nitrogen in bone collagen. Nitrogen isotope ratios of bone collagen show that Jinggouzi ancient population in primarily ate animal products with only a small amount of plant products. Carbon isotope ratios of bone collagen show that most plant products come from C₄ plant.

Physical anthropology

Obtaining population genetics data via non-destructive means: a three-dimensional analysis of human craniofacial morphology

Sabrina B. Sholts1, Sebastian Wärmländer2, & Phillip L. Walker3
1 Department of Anthropology, University of California at Santa Barbara, Department of Anthropology, University of California, Santa Barbara, CA 93106, USA

2 Division of Biophysics, Arrhenius Laboratories for Natural Science, Stockholm University, 10691 Stockholm, Sweden

3 Department of Anthropology, University of California at Santa Barbara, Department of Anthropology, University of California, Santa Barbara, CA 93106, USA

In this study, shapes extracted from the human craniofacial skeleton were used to investigate the genetic heritabilities of morphological traits. Three-dimensional point data was collected from human crania with a 3D laser scanner and used to render complete 3D surface models of the original skeletal material. 3D data analysis software was used to digitally slice the cranial models with geometric planes defined by traditional craniometric landmarks. The cross-sections produced by these planes yielded contours of cranial outlines for different craniofacial features. Using elliptical Fourier transforms, the contours were parametrized into series of Fourier coefficients, which, due to their inherent orthogonality, form suitable input parameters for statistical analysis. Principal components analysis (PCA) was employed to differentiate population groups based on shape differences in various aspects of cranio-facial skeletal morphology. This method can yield population genetics data and information on probable ancestral affinity using non-destructive analysis of human remains and with greater accuracy than with traditional craniometric studies of metric and non-metric traits. The application of this method to human skeletal collections can elucidate genetic relationships in past populations and improve our understanding of their archaeological contexts. This non-invasive method also offers a viable alternative for determining ancestral affinities between groups and individuals in cases where DNA testing is not possible, due to either the necessary destruction of bone required for DNA analysis or the degraded condition of the material.