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Morris more nodding jibbing shoots wonderfully? By integrating the molecular data A negative control was included in every PCR round. Electropherogram analysis and sequence New DNA sequences of modern bears from the Iberian assembly was conducted in Geneious 7. Peninsula com. During electropherogram analysis, the primer annealing In order to study the genetic legacy of the Pleistocene and ancient regions and the low quality regions at both ends of each elec- Holocene bears in the Iberian Peninsula we increased the data- tropherogram were trimmed error probability limit of 0.
The other modern sample individual Ua come DNA sequences of bears belonging to subclade 1a that covered from hairs from a skin seized from poachers. The museum sam- a wide geographic range, including the Italian, Iberian and ples were hair of brown bear from the Pyrenees preceding the Scandinavian Peninsulas, southeast France and the British Isles introduction of Eastern European bears in the area, donated by Supplemental online material, Table S We aligned published sequences to the ones generated in this Extraction and sequencing of modern biological samples was study, compiling a data-set of bears of subclade 1a Table S For Modern samples sequences convenience of the study, 6 sequences from subclade 1b Taberlet and Bouvet ; Hofreiter et al.
The longest sequences Haplotypes were determined by DnaSP v5 Librado and Rozas belong to the modern Cantabrian haplotype, being all of them The sequences reported in this For temporal analysis of subclade 1a, a network has been made paper have been deposited in the GenBank database accession using TempNet, an R-script Prost and Anderson MF — MF Mitochondrial haplotypes Table 1. F-1 Somiedo Can- — CR MF Scandinavian and British brown bears form up the central Asturi- tabri- part of the network, with the haplotype 15 being the central one.
An exhaustive survey in the literature produced a data-set of The same occurs with the scarce representation of mainland direct radiocarbon ages of Western European brown bears and Pleistocene brown bears. Based on the Based on the 14C dates and the geographic origin of the sam- genetic information available, clade 1b seems to have dis- ples, towards the end of the Pleistocene and before the LGM, appeared from the British Isles after the LGM, as all fossils we observe the presence of brown bears in the British Isles analysed so far belongs either to clade 1a or to clade 2. Most mainly in Ireland , Continental Western Europe and the Iberian of the dated subclade 1a bears are concentrated in the very Peninsula.
Furthermore, two In the Iberian Peninsula it seems to be a gap in the fossil Pleistocene individuals from the Iberian Peninsula were found record around the LGM, as only one sample has been dated to to belong to clades 3c Valdiosera et al. Interestingly, all Iberian Jacobi and Higham ; Edwards et al. Downloaded by [University of Coruna] at 04 October Figure 3. Notes: Clade 1 — circle, clade 2 — diamond, clade 3 — inverted triangle, clade 4 — triangle. For more information, see supplementary Table S In order to investigate the temporal and geographical distribu- About other areas in Western Europe, there is a general lack of tion of the maternal lineages of brown bear in Western Europe, fossil information.
Something similar happens in France and other The Figure 4 clearly shows that the Pleistocene haplotypes of areas from Continental Western Europe, where most of the fos- the Iberian Peninsula have no continuity in MIS 1.
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Network of subclade 1a from MIS 3 to nowadays. Notes: Modern haplotypes are in bold line. MIS 1 and is found in historical times in Southern Scandinavia history of brown bear diversification in this area of northern Taberlet and Bouvet ; Xenikoudakis et al. However, there is no indication that the Iberian bears, is greatly diversified.
Sub fossil or ing peripheral positions in the network, clearly distant from the historical individuals nineteenth and twentieth century show Iberian Pleistocene bears and from haplotype 15, the central one more genetic diversity, sharing two haplotypes with the Britain in the network and continuous along the post-glacial stage. There are only 2 direct dated fossils from south Scandinavia and one indirectly dated from Denmark.
These bears record the expansion been analysed at genetic level belong to the subclades 1b and 2. However, there is not genetic data from reappeared in the Holocene in Ireland Edwards et al. The Irish clade 2 bears probably did not survive in a proposed The colonization of Scandinavia by Clade 1a brown bears refugium in southwest Ireland Leonard et al.
The Montgomery et al.
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The lithostatic discharge Sea shelf, from where they re-entered Ireland when suitable areas caused by the gradual melting of the ice cap caused lithosphere Downloaded by [University of Coruna] at 04 October were available. If these bears were hybrids of brown and polar bears, Borns ; Figure 2—6. In the absence of data from bears, they could have found suitable habitats in these coastal areas. An extensive sea ice cover would However, remnants of ice sheet were still isolating the south have flanked these ice sheets, and many of the small islands pres- and the north of the Scandinavian Peninsula.
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This barrier ent in the Celtic Sea are likely to have become connected Brooks must have prevented subclade 1a bears to expand to north- et al. In such a scenario it would not be rare that clade 2 ern Scandinavia for millennia. Later, when the ice disappeared brown bears could have reached southern latitudes during the Andersen and Borns ; Figure 6. We have not found genetic data of brown bears already colonized by bears of clade 3 from Eastern Europe. The in the Bay of Biscay area Aquitaine, Basque Country , but the suture zone between both clades is still present Taberlet et al.
Another possi- As mentioned previously, among the 8 Iberian Pleistocene bears ble polar bear representation comes from Isturitz cave in the with genetic data there is no putative ancestor of the bears that northern flank of the Pyrenees Atlantic Pyrenees, Aquitaine. Furthermore, rib showing two individuals that somehow resemble polar bears two Pleistocene individuals from the Iberian Peninsula were Esparza San Juan and Mujika Alustiza , Figure 1.
No other repre- 1a. In fact, according to radiocarbon dates, Britain was the first sentatives of these clades in the Iberian Peninsula or in Western area occupied by this subclade after the LGM. The origin of Europe are known, neither fossil nor alive. Clade 3c has only been those bears is unknown; if they were traditionally supposed to found in fossil bears from North America Barnes et al.
United States in North America, and the nearest members of Barriers such as the Pyrenees or the Fleuve Manche would had this clade in Europe are historical individuals from Caucasus acted as a barrier for the expansion of the Iberian bears, as sug- Bray Based Scandinavia on the genetic information available from fossil and modern Modern populations from southern Scandinavia show only two individuals, it seems that these two individuals represent a minor haplotypes, one in the Norwegian population, and another in the population of vagrant bears that expanded westwards, instead Swedish one Taberlet and Bouvet ; Kohn et al.
The effective population size of both lineages would for several months, so they are more prone to disturbance by have been small, since finally they faded out without contributing human groups Swenson et al. According to this hypothe- populations nor in the present day Iberian bears. In fact, the Barton ; Edwards et al. More to the northwest, in respecting the early recolonization of the British Isles. Throughout the Holocene, the record of brown bears in the N We could argue that this hiatus is an artefact caused by the of the Iberian Peninsula become scarcer, probably in response to almost total absence of direct dated samples in the many sites of the anthropic pressure culminating in the almost total extinction Cantabria and the Basque Country, that is the natural entrance of the species.
Brown bears began to reduce their population of the North Corridor. In fact, the intense archaeological studies drastically since the appearance of firearms in sixteenth century, in the East Cantabrian area yielded up to 6 sites with brown bear from south to north. Between 0. However, up to 40 sites Nores and Naves with a total of 40 individuals Couturier in the same area did not yield any Solutrean or Magdalenian bear In that decade, Naves and Palomero describe a remain, but related to older or younger archaeological contexts.
While it would be desirable to have more direct dated samples All Cantabrian modern bears the oldest living in the s, from this region, it does not seem possible that hypothetic early Valdiosera et al. This haplotype is missing in the Holocene samples. This is the result of a strong bottleneck caused by the anthropic This late entry has several possible explanations.
In addition, the modern haplotype was just not found in the Holocene samples, proximity of the Atlantic Pyrenees to the coast means a bar- or if it arrived more recently from the central or south Iberian rier for the bears to entry in the peninsula. Studies on glacial Peninsula. Jalut et al.
Several phases of advance and retreat of glaciers in the Iberian Peninsula.