Ancient DNA Reveals Matrilineal Continuity in Present-Day Poland over the Last Two Millennia

While numerous ancient human DNA datasets from across Europe have been published till date, modern-day Poland in particular, remains uninvestigated. Besides application in the reconstruction of continent-wide human history, data from this region would also contribute towards our understanding of the history of the Slavs, whose origin is hypothesized to be in East or Central Europe. Here, we present the first population-scale ancient human DNA study from the region of modern-day Poland by establishing mitochondrial DNA profiles for 23 samples dated to 200 BC – 500 AD (Roman Iron Age) and for 20 samples dated to 1000–1400 AD (Medieval Age). Our results show that mitochondrial DNA sequences from both periods belong to haplogroups that are characteristic of contemporary West Eurasia. Haplotype sharing analysis indicates that majority of the ancient haplotypes are widespread in some modern Europeans, including Poles. Notably, the Roman Iron Age samples share more rare haplotypes with Central and Northeast Europeans, whereas the Medieval Age samples share more rare haplotypes with East-Central and South-East Europeans, primarily Slavic populations. Our data demonstrates genetic continuity of certain matrilineages (H5a1 and N1a1a2) in the area of present-day Poland from at least the Roman Iron Age until present. As such, the maternal gene pool of present-day Poles, Czechs and Slovaks, categorized as Western Slavs, is likely to have descended from inhabitants of East-Central Europe during the Roman Iron Age.     

Cultivation and processing of Linum usitatissimum and Camelina sativa in southern Scandinavia during the Roman Iron Age

Charred seed conglomerates of Linum usitatissimum (flax) and Camelina sativa (gold of pleasure) were found at Uppåkra 2:25, a Roman Iron Age site in Skåne, southern Sweden. The conglomerates showed no mixing with each other, as they were almost pure flax and gold of pleasure respectively. Together with other archaeobotanical data from the site, they provide new evidence on the use, processing and cultivation of these two plants in early Iron Age in Scandinavia. Metric analyses were applied to flax seeds from both conglomerates and other contexts at this site, and compared to seed assemblages from other Roman Iron Age sites in Europe. The comparison showed that the flax cultivation at Uppåkra 2:25 was intended for the production of oil-rich seeds. The contextual relationship indicates that both flax and gold of pleasure seeds were processed in a similar way and used for oil. Furthermore, the pure seed conglomerate of gold of pleasure suggests that this plant was not a weed, but rather an intentionally grown crop which was cultivated separately from flax.     

Rhytidium rugosum (Bryophyta) colonized Scandinavia from at least two glacial refugial source populations

The Scandinavian post‐glacial history of the moss Rhytidium rugosum is traced on the basis of information from the nuclear markers ITS and gpd for 229 Scandinavian and 81 other specimens. Some haplotypes, groups or lineages identified in a NeighborNet split network are predominantly northern Scandinavian, whereas others are southern. With the distributions of individual haplotypes and the timing of the deglaciation in different parts of Scandinavia, this implies colonization from the south and from the north or north‐east. High haplotype and nucleotide diversity and the occurrence of certain private haplotypes in the north suggest that the species may have survived the Last Glacial Maximum in local refugia. Slightly higher numbers of private haplotypes in Scandinavia than in central or north‐eastern Europe also favour an explanation with at least some local glacial survival. Low diversity in the southernmost contiguous region of the Scandinavian mountain range is probably a result of recent land uplift and late colonization. The Scandinavian lowland regional populations probably represent remains of an earlier widespread population that became increasingly restricted to small and isolated areas when the vegetation closed during the post‐glacial period. Some of the lowland populations require extensive management to survive. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 635–657.     

Genetic Diversity among Ancient Nordic Populations

Using established criteria for work with fossil DNA we have analysed mitochondrial DNA from 92 individuals from 18 locations in Denmark ranging in time from the Mesolithic to the Medieval Age. Unequivocal assignment of mtDNA haplotypes was possible for 56 of the ancient individuals; however, the success rate varied substantially between sites; the highest rates were obtained with untouched, freshly excavated material, whereas heavy handling, archeological preservation and storage for many years influenced the ability to obtain authentic endogenic DNA. While the nucleotide diversity at two locations was similar to that among extant Danes, the diversity at four sites was considerably higher. This supports previous observations for ancient Britons. The overall occurrence of haplogroups did not deviate from extant Scandinavians, however, haplogroup I was significantly more frequent among the ancient Danes (average 13%) than among extant Danes and Scandinavians (∼2.5%) as well as among other ancient population samples reported. Haplogroup I could therefore have been an ancient Southern Scandinavian type “diluted” by later immigration events. Interestingly, the two Neolithic samples (4,200 YBP, Bell Beaker culture) that were typed were haplogroup U4 and U5a, respectively, and the single Bronze Age sample (3,300–3,500 YBP) was haplogroup U4. These two haplogroups have been associated with the Mesolithic populations of Central and Northern Europe. Therefore, at least for Southern Scandinavia, our findings do not support a possible replacement of a haplogroup U dominated hunter-gatherer population by a more haplogroup diverse Neolithic Culture.     

Uniform diet in a diverse society. Revealing new dietary evidence of the Danish Roman Iron Age based on stable isotope analysis

A systematic dietary investigation during Danish Roman Iron Age (1‐375AD) is conducted by analyzing stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) in the collagen of human and animal bone. The human sample comprises 77 individuals from 10 burial sites. In addition 31 samples of mammals and fish were analyzed from same geographical area. The investigation characterizes the human diet among different social groupings and analyses dietary differences present between sex, age, and site phase groups. Diachronically, the study investigates the Roman influences that had an effect on social structure and subsistence economy in both the Early and Late Period. Geographically the locations are both inland and coastal. The isotopic data indicate extremely uniform diet both between and within population groups from Early and Late Roman periods and the data are consistent throughout the Roman Iron Age. Protein consumption was dominated by terrestrial animals with no differences among social status, age, sex, or time period, while terrestrial plant protein only seems to have contributed little in the diet. Furthermore, the consumption of marine or aquatic resources does not seem to have been important, even among the individuals living next to the coast. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Late Holocene landscape development around a Roman Iron Age mass grave,Alken Enge,Denmark

Sediments from the small lake Ilsø situated in the Illerup/Alken Enge Valley were studied in order to investigate past landscape development at the time of a probably ritual human mass burial following battle during the Roman Iron Age (ad 1–400). A pollen record from Ilsø and a number of other records from Jutland were combined using the Landscape Reconstruction Algorithm to reconstruct local vegetation changes through the last 2,800 years. These methods were supplemented by studies of catchment-related geochemistry of the Ilsø lake sediments. The results show a marked reforestation event associated with a strong decrease in erosion levels at the very beginning of the first century ad, contemporaneous with the finds of human remains at Alken Enge. Comparison with a pollen record 10 km away and with those from other sites, reveals that this reforestation occurs unusually early and rapidly, and is an unparalleled development in a Danish context. We suggest that the major landscape changes at the beginning of the Roman Iron Age and forest cover for the next few centuries comprise a possible example of ritual control of local land-use.     

mtDNA analysis of human remains from an early Danish Christian cemetery

One of Denmark's earliest Christian cemeteries is Kongemarken, dating to around AD 1000-1250. A feature of early Scandinavian Christian cemeteries is sex segregation, with females buried on the northern sides and males on the southern sides. However, such separation was never complete; in the few early Christian cemeteries excavated in Scandinavia, there were always a few males placed on the north side, and some females on the south side. At Kongemarken, several males with juxtaposed females were found on the north side of the cemetery. Thus, to evaluate possible kinship relationships, and more general questions of population affinity, we analyzed mitochondrial DNA extracted from nine individuals excavated in two different areas within the cemetery: one male and four females from Area 1, and one male and three females from Area 2. Using stringent laboratory protocols, each individual was unequivocally assigned to a mitochondrial haplogroup. A surprising amount of haplogroup diversity was observed (Area 1: 1 U7 (male), 1 H, 1 I, 1 J, and 1 T2; Area 2: 2 H, 1 I, and 1 T, with one H being male); even the three subjects of haplogroup H were of different subtypes. This indicates that no subjects within each area were maternally related. The observed haplogroup, U7, while common in India and in western Siberian tribes, was not previously observed among present-day ethnic Scandinavians, and haplogroup I is rare (2%) in Scandinavia. These observations suggest that the individuals living in the Roskilde region 1,000 years ago were not all members of a tightly knit local population and comprised individuals with genetic links with populations that were from much farther away.     

The history of rye cultivation in Europe

During recent years finds from several prehistoric and medieval periods have thrown new light on the history of the spread of rye. It is now proven that wild rye is indigenous to Anatolia and was already domesticated there by the early Neolithic at the beginning of agriculture. Secale migrated to Central Europe as a weed among other cereals, and single grains of weed rye have been recorded there since the early Neolithic. The number of finds increased during the Bronze Age and Iron Age, and the status of rye changed from weed to crop plant, probably in the course of the early Iron Age. This acculturation of Secale cereale in central and eastern Europe was obviously independent of the earlier one in Anatolia. The first stages towards deliberate cultivation happened unintentionally through harvesting close to the ground, so that the rye was permanently represented in the seed corn. From this point rye was able to take advantage of its competitive strength on poor soils and in areas with unfavourable climate. The start of rye as a crop in its own right during the pre-Roman Iron Age and Roman period presumably took place independently in different areas. The expansion of intensive rye cultivation occurred in the Middle Ages. However, new finds from north-west Germany, which are presented here, show that in this area rye has been cultivated as a main crop on poor soils since the Roman period. In two maps all rye finds up to 1000 A.D. are shown, which after critical consideration can be regarded as cultivated rye.     

"These boots were made for walking": the isotopic analysis of a C(4) Roman inhumation from Gravesend, Kent, UK

As part of the road widening scheme between London and Dover, Oxford Archaeology South uncovered a large boundary ditch of Iron Age origin that contained Iron Age and Roman inhumations, adjacent to which was a small mid-late Roman cemetery, interpreted as a rural cemetery for Romano-British farmers. Grave goods in the cemetery were restricted to a few individuals with hobnailed boots. Bulk bone collagen isotopic analysis of 11 skeletons of Iron Age and Roman date gave a typical C(3) terrestrial signal (average δ(13) C = -19.8‰, δ(15) N = 9.3‰), but also revealed one (SK12671) with a diet which included a substantial C(4) component (δ(13) C = -15.2‰, δ(15) N = 11.2‰). This is only the second such diet reported in Roman Britain. Subsequent δ(18) O(c) and (87) Sr/(86) Sr measurements on the dental enamel in this individual were, however, consistent with a "local" origin, indicating that either C(4) protein was consumed in Late Roman Britain, or that he came from somewhere else, but where conditions gave rise to similar isotopic values. If we accept the latter, then it indicates that using oxygen and strontium isotopes alone to identify "incomers" may be problematic. The provision of hobnailed boots for the dead appears to have had a strong symbolic element in Late Roman Britain. We suggest that in this case the boots may be significant, in that he was being equipped for the long march home.     

Adaptive evolution signals in mitochondrial genes of Europeans

Since modern Europeans appear to be descendants of the Late Pleistocene European peoples who survived the last glacial period, it is quite reasonable to expect the presence of adaptive genetic variants that originated in the Ice Age in the modern gene pool of Europeans. To find such adaptive variants, mitochondrial genomes have been analyzed of the modern population from Eastern and Central Europe belonging to haplogroups U4, U5, and V, that diversified during the Late Pleistocene and Holocene periods. Analysis of distribution of nonsynonymous and synonymous substitutions, as well as results of search for radical amino acid changes that arose under the influence of adaptation (positive destabilizing selection) allowed us to detect signals of molecular adaptation in different mitochondrial genes and haplogroups of mtDNA. However, there were very few strong adaptive signals (z > 3.09, P < 0.001) that could be due to the loss of adaptive mtDNA haplotypes during the Holocene warming.     

Maternal and paternal genetic diversity of ancient sheep in Estonia from the Late Bronze Age to the post‐medieval period and comparison with other regions in Eurasia

Sheep were among the first domesticated animals to appear in Estonia in the late Neolithic and became one of the most widespread livestock species in the region from the Late Bronze Age onwards. However, the origin and historical expansion of local sheep populations in Estonia remain poorly understood. Here, we analysed fragments of the hypervariable D‐loop of mitochondrial DNA (mtDNA; 213 bp) and the Y‐chromosome SRY gene (130 bp) extracted from 31 archaeological sheep bones dated from approximately 800 BC to 1700 AD. The ancient DNA data of sheep from Estonia were compared with ancient sheep from Finland as well as a set of contemporary sheep breeds from across Eurasia in order to place them in a wider phylogeographical context. The analysis shows that: (i) 24 successfully amplified and analysed mtDNA sequences of ancient sheep cluster into two haplogroups, A and B, of which B is predominant; (ii) four of the ancient mtDNA haplotypes are novel; (iii) higher mtDNA haplotype diversity occurred during the Middle Ages as compared to other periods, a fact concordant with the historical context of expanding international trade during the Middle Ages; (iv) the proportion of rarer haplotypes declined during the expansion of sheep from the Near Eastern domestication centre to the northern European region; (v) three male samples showed the presence of the characteristic northern European haplotype, SNP G‐oY1 of the Y‐chromosome, and represent the earliest occurrence of this haplotype. Our results provide the first insight into the genetic diversity and phylogeographical background of ancient sheep in Estonia and provide basis for further studies on the temporal fluctuations of ancient sheep populations.     

Infectious Pancreatic Necrosis First Isolation of Virus from Fish in Norway

Infectious pancreatic necrosis (IPN) is a disease of salmonid fishes. It has been reported in many countries throughout the world (M’Gonigle 1940, Wood et al. 1955, Besse & Kinkelin 1965, Vestergård Jørgensen & Bregnballe 1969, Sano 1971, Ball et al 1971, Ljungberg & Vestergård Jørgensen 1972, Schlotfeldt et al. 1975). Outbreaks of the disease can cause serious losses in populations of hatchery reared salmonids, the mortality rate varying between 10 and 90 % (Vestergård Jørgensen & Kehlet 1971). There are at least four different serotypes of the virus distinguished by neutralization tests (Wolf et al. 1968). Twenty-five isolates of IPN virus in Denmark proved to represent only two serotypes (Sp and Ab) (Vestergård Jørgensen & Kehlet). The present paper reports the first isolation of IPN virus from the stock at a fish farm in Norway.     

A new approach to the study of Romanization in Britain: a regional perspective of cultural change in late iron age and roman dorset using the siler and gompertz-makeham models of mortality

This is the first study of health in the Roman Empire to use the Siler and Gompertz-Makeham models of mortality to investigate the health consequences of the 43 AD conquest of Britain. The study examined late Iron Age and Romano-British populations (N = 518) from Dorset, England, which is the only region of Britain to display continuity in inhumation burial practice and cemetery use throughout the two periods. Skeletal evidence for frailty was assessed using cribra orbitalia, porotic hyperostosis, periosteal lesions, enamel hypoplasia, dental caries, tuberculosis, and rickets. These health variables were chosen for analysis because they are reliable indicators of general health for diachronic comparison (Steckel and Rose: The backbone of history: health and nutrition in the western hemisphere (2002)) and are associated with the introduction of urbanism in Britain during the Roman period (Redfern: J Rom Archaeol Supp Series 64 (2007) 171-194; Redfern: Britannia 39 (2008a) 161-191; Roberts and Cox: Health and disease in Britain: from prehistory to the present day (2003)). The results show that levels of frailty and mortality were lower in the late Iron Age period, and no sex differences in mortality was present. However, post-conquest, mortality risk increased for children and the elderly, and particularly for men. The latter finding challenges received wisdom concerning the benefits of incorporation into the Empire and the higher status of the male body in the Roman world. Therefore, we conclude that the consequences of urbanism, changes in diet, and increased population heterogeneity negatively impacted health, to the extent that the enhanced cultural buffering of men did not outweigh underlying sex differences in biology that advantage women.     

Population history of the middle Euphrates valley: Dental non-metric traits at Tell Ashara,Tell Masaikh and Jebel Mashtale,Syria

Fifty-nine dental non-metric traits were scored using Arizona State University Dental Anthropology System on a sample of teeth from 350 human skeletons excavated at three sites in the lower middle Euphrates valley. The dataset was divided into six chronological subsets: Early Bronze Age, Middle Bronze Age, Early Iron Age with Neo-Assyrian period, Classical/Late Antiquity, Early Islamic (Umayyad and Abbasid) period and Modern period. The matrix of Mean Measure of Divergence values exhibited temporal homogeneity of the sample with only dental non-metric trait scores in the Modern subset differing significantly from most other subsets. Such a result suggests that no major gene flow occurred in the middle Euphrates valley between the 3rd millennium BCE and the early 2nd millennium CE. Only after the Mongolian invasion and large depopulation of northern Mesopotamia in the 13th century CE a major population change occurred when the area was taken over in the 17th century by Bedouin tribes from the Arabian Peninsula.     

Human impact signals from peat bogs – a combined palynological and geochemical approach

This research compares palynological evidence for changes in land use histories with a geochemical method for reconstructing past soil erosion. Changes in land use have significant effects on soil erosion. It has been shown elsewhere that silicon (Si) and titanium (Ti) are good proxies for soil erosion. Ombrotrophic peat bogs are useful archives in which to measure Si and Ti depositions as they only receive inorganic erosional inputs through atmospheric deposition and they contain very low background levels of mineral matter. The correlation between geochemical and pollen analytical reconstructions of past human activity from three raised bog sites in Great Britain and Ireland is discussed here, with reference to examples from four particular time periods: the mid-to-late Bronze Age/Iron Age, the late Iron Age/Roman period, the Middle Ages/Tudor period and the more recent past. The results generally indicate a close correlation between the palynological and geochemical proxies, with the combination of both methods allowing a more comprehensive interpretation of the palaeoenvironmental record. Plantago lanceolata and Poaceae pollen frequencies appear to correlate particularly well with the geochemical proxies. A multi-proxy approach such as this may be particularly useful for identifying and interpreting low-level prehistoric human impact.     

The life cycle of Cyclops vicinus in Lake Søbygård: new aspects derived from sediment analyses

Cyclops vicinus is the only copepod species in the pelagic zone of Lake Søbygård and can be found there all year round. We studied the population dynamics of this copepod over a one-year period. In contrast to earlier studies we included the copepods in the pelagic zone as well as the copepods resting in the sediment. Cyclops vicinuswas found not only in the open water, but also in the sediment throughout the year. From the fluctuations of the abundances in both habitats we suggest that the life cycle is more complex than known so far: different diapause pattern appear within the population and the induction of diapause is different than in other populations. We assume that these features contribute to the predominance of C. vicinusin Lake Søbygård.     

Genetic differentiation in Hippocrepis emerus (Leguminosae): allozyme and DNA fingerprint variation in disjunct Scandinavian populations

The structure of genetic variation in disjunct Scandinavian populations of Hippocrepis emerus was studied using allozymes and DNA fingerprinting. Variation in the three native regional populations in Scandinavia was compared with that in a recently introduced population in Sweden. In contrast to the recently introduced population, the native Scandinavian isolates of H. emerus showed high levels of allozyme fixation and low levels of DNA diversity. Variation in allozymes and at DNA fingerprint loci showed closely congruent patterns of geographic variation, with pronounced differentiation between the native Norwegian and Swedish isolates of the species. The structure of genetic variation in native Scandinavian H. emerus is interpreted in terms of historical population bottlenecks and founder events during the species' postglacial immigration into Scandinavia.     

Multilocus phylogeography and population structure of common eiders breeding in North America and Scandinavia

Aim Glacial refugia during the Pleistocene had major impacts on the levels and spatial apportionment of genetic diversity of species in northern latitude ecosystems. We characterized patterns of population subdivision, and tested hypotheses associated with locations of potential Pleistocene refugia and the relative contribution of these refugia to the post‐glacial colonization of North America and Scandinavia by common eiders (Somateria mollissima). Specifically, we evaluated localities hypothesized as ice‐free areas or glacial refugia for other Arctic vertebrates, including Beringia, the High Arctic Canadian Archipelago, Newfoundland Bank, Spitsbergen Bank and north‐west Norway. Location Alaska, Canada, Norway and Sweden. Methods Molecular data from 12 microsatellite loci, the mitochondrial DNA (mtDNA) control region, and two nuclear introns were collected and analysed for 15 populations of common eiders (n = 716) breeding throughout North America and Scandinavia. Population genetic structure, historical population fluctuations and gene flow were inferred using F‐statistics, analyses of molecular variance, and multilocus coalescent analyses. Results Significant inter‐population variation in allelic and haplotypic frequencies were observed (nuclear DNA F_ST = 0.004–0.290; mtDNA Φ_ST = 0.051–0.927). Whereas spatial differentiation in nuclear genes was concordant with subspecific designations, geographic proximity was more predictive of inter‐population variance in mitochondrial DNA haplotype frequency. Inferences of historical population demography were consistent with restriction of common eiders to four geographic areas during the Last Glacial Maximum: Belcher Islands, Newfoundland Bank, northern Alaska and Svalbard. Three of these areas coincide with previously identified glacial refugia: Newfoundland Bank, Beringia and Spitsbergen Bank. Gene‐flow and clustering analyses indicated that the Beringian refugium contributed little to common eider post‐glacial colonization of North America, whereas Canadian, Scandinavian and southern Alaskan post‐glacial colonization is likely to have occurred in a stepwise fashion from the same glacial refugium. Main conclusions Concordance of proposed glacial refugia used by common eiders and other Arctic species indicates that Arctic and subarctic refugia were important reservoirs of genetic diversity during the Pleistocene. Furthermore, suture zones identified at MacKenzie River, western Alaska/Aleutians and Scandinavia coincide with those identified for other Arctic vertebrates, suggesting that these regions were strong geographic barriers limiting dispersal from Pleistocene refugia.     

Ancient DNA reveals a migration of the ancient Di‐qiang populations into Xinjiang as early as the early Bronze Age

Xinjiang is at the crossroads between East and West Eurasia, and it harbors a relatively complex genetic history. In order to better understand the population movements and interactions in this region, mitochondrial and Y chromosome analyses on 40 ancient human remains from the Tianshanbeilu site in eastern Xinjiang were performed. Twenty‐nine samples were successfully assigned to specific mtDNA haplogroups, including the west Eurasian maternal lineages of U and W and the east Eurasian maternal lineages of A, C, D, F, G, Z, M7, and M10. In the male samples, two Y chromosome haplogroups, C* and N1 (xN1a, N1c), were successfully assigned. Our mitochondrial and Y‐chromosomal DNA analyses combined with the archaeological studies revealed that the Di‐qiang populations from the Hexi Corridor had migrated to eastern Xinjiang and admixed with the Eurasian steppe populations in the early Bronze Age. Am J Phys Anthropol 157:71–80, 2015. © 2014 Wiley Periodicals, Inc.     

Ancient DNA identifies post‐glacial recolonisation,not recent bottlenecks,as the primary driver of contemporary mtDNA phylogeography and diversity in Scandinavian brown bears

Aim

Brown bear populations in Scandinavia show a strong mitochondrial DNA (mtDNA) phylogeographic structure and low diversity relative to other parts of Europe. Identifying the timing and origins of this mtDNA structure is important for conservation programs aimed at restoring populations to a natural state. Therefore, it is essential to identify whether contemporary genetic structure is linked to post‐glacial recolonisation from divergent source populations or an artefact of demographic impacts during recent population bottlenecks. We employed ancient DNA techniques to investigate the timing and potential causes of these patterns.

Location

Scandinavia and Europe.

Methods

Ancient mtDNA sequences from 20 post‐glacial Scandinavian bears were used to investigate phylogeographic structure and genetic diversity over the last 6000 years. MtDNA from 19 Holocene Norwegian bears was compared with 499 sequences from proximate extant populations in Sweden, Finland, Estonia and western Russia. A single mtDNA sequence from a Holocene Denmark sample was compared with 149 ancient and modern bears from Western Europe.

Results

All nineteen Holocene Norwegian samples are identical to or closely related to the most common mtDNA haplotype found in northern Europe today. MtDNA diversity was low and not significantly different from extant populations in northern Europe. In Denmark, we identified a single mtDNA haplotype that is previously unrecorded from Scandinavia.

Main conclusions

The current discrete phylogeographic structure and lack of mtDNA diversity in Scandinavia is attributed to serial founder effects during post‐glacial recolonisation from divergent source populations rather than an artefact of recent anthropogenic impacts. In contrast to previous interpretations, the recolonisation of southern Scandinavia may not have been limited to bears from a single glacial refugium. Results highlight the importance of conserving the long‐term evolutionary separation between northern and southern populations and identify southern Scandinavia as an important reservoir of mtDNA diversity that is under threat in other parts of Europe.