What makes up iberian peninsula




















Some of the native species include Pyrenean desman, shrew, and European hedgehog. Other species include Eurasian beaver, Alpine marmot, the European rabbit, Iberian lynx, Italian wolf, and Eurasian lynx.

Humans have been living on the Iberian Peninsula since over 1. Neanderthals occupied the peninsula around , BP. However, early modern humans migrated to the peninsula approximately 40, years ago from Southern France. The Roman army occupied the peninsula around BC after successfully fighting the Carthaginians. However, it took Rome years to defeat the Iberians and Celtic and annex the peninsula. Thereafter, Hispania Province was established.

The Romans occupied the Iberian Peninsula for years. The Germanic people, including Vandals, Suebi, and Alans began settling on the peninsula around the 5th century.

The Visigoths arrived much later and occupied the whole of the peninsula, expelling other tribes except the Suebi people. However, around , they took control of the Suebi Kingdom, including Bracara Braga. In the early 8th century, Muslims, led by Tariq Ibn Ziyad conquered and occupied the peninsula except for the northern kingdom. By the end of Muslim dominance on the peninsula end of 15th century , the Iberian Peninsula had an estimated population of about 6.

The Iberian Peninsula has an estimated population of 53 million people and is dominated by three main metropolises; Lisbon, Barcelona, and Madrid. Madrid is the largest metropolitan region, with a population of 6. Barcelona has a population of about 5. Valencia and Seville in Spain have a combined population of 4. These countries have some of the thriving industries in Europe, including fishing, tourism, and mining.

Because of the long coastline, fishing is a popular economic activity, with major catches including tuna and sardines. Processed fish products from Spain and Portugal are exported to the global market through several companies, including Ramirez. Besides, it also produces barley, corn, and wheat. The other countries part of this region are Portugal , France , Andorra , and Gibraltar. Several major cities are located on the Iberian Peninsula.

The largest is Madrid , Spain, which has a population of over 3. Barcelona , which is also located in Spain, is the second most populous city in the Iberian Peninsula with a population of over 1. Iberian Peninsula Countries Fuente Pecina 1, Fuente Pecina 2, Fuente Pecina 4, Alto de Reinoso, La Mina, La Tarayuela, El Juncal, Arroyal I, El Hundido, Camino de las Yeseras, Humanejos, Valle de las Higueras, 60— El Mirador, Es Forat de ses Aritges.

With regard to its population history, the Iberian Peninsula has been the focus of several recent archaeogenetic studies 39 , 40 , 41 , 42 , 43 , Studies focusing not only on mtDNA but also on Y chromosome markers have supported the model of a pioneer colonization in the north-eastern coastal regions of the Iberian Peninsula at the onset of the Iberian Neolithic 39 , On the other hand, the northern part of Spain Cantabrian fringe attested to a rather complex Neolithic transition Recent mtDNA and genome-wide analyses have put an emphasis on the genetic affinity and shared Near Eastern ancestry between the early Iberian farmers and the contemporary Central European Linearbandkeramik LBK population 46 , 47 , The latest mtDNA and genomic studies have revealed increased subsequent admixture of hunter-gatherer elements during the local middle Neolithic La Mina; Alto de Reinoso, n.

Despite numerous research projects being carried out over the past years, the Neolithic settlement history of Europe can still only be explained at a broad scale 47 , 49 , 51 , 52 , 53 , Regional transects through time detailing the developments and the course of the Neolithic in central Germany 55 , 56 , 57 and in the Carpathian Basin 58 have mostly been examined by mitochondrial DNA mtDNA control region data. Our project completes the latter series by focusing on the archaeological models and hypotheses that have been put forward for the Iberian Peninsula, and where diachronic i.

Essential research questions focus on three levels: i the individual sites, ii the Iberian Peninsula as a scene of Neolithic transition and iii a comparison with contemporaneous ancient and modern-day Europeans. A key question of our study of the mtDNA diversity on the Iberian Peninsula through time was to examine to what extent regional and supra-regional cultural groups could be recognized as genetically identifiable entities, as shown in other areas of Europe 46 , A related question was whether the cultural breaks that can be seen, for instance, at the end of the Neolithic and Chalcolithic periods, were also accompanied by human population turnovers.

We processed ancient DNA samples of human individuals from prehistoric Iberia and generated reproducible mitochondrial hypervariable region haplotypes HVS-I, np — from individuals following strict authentication criteria see Methods.

The resulting haplogroup compositions of these groups are presented in Fig. MtDNA haplogroup composition of the prehistoric Iberian groups. Relative haplogroup frequencies are presented in Supplementary Table S6.

Recent NGS studies described hunter-gatherers belonging to haplogroup U5b in north Iberia 43 , 59 , while the suggested pre-farming presence of haplogroups H, N and U4 42 were reported in studies that did not meet our strict authentication criteria see Methods. We used the haplogroup frequencies of the studied groups for principal component analyses PCA, see Methods.

S1—2 , but it also shows some affinities with further Iberian farmers through common EN haplogroups e.

K, J, T2 Haplogroup compositions of all other Iberian groups are highly similar to each other, resulting in clusters on the PCA plots and Ward clustering tree. Principal component analysis based on haplogroup frequencies of individuals from 16 prehistoric groups. The first two components display For abbreviations of Iberian groups, see legend of Fig. For further information see Supplementary Table S6. The combined dataset shows a predominance of haplogroup K An interesting exception is haplogroup L1b in the Late Chalcolithic central Iberia at the site Camino de las Yeseras n.

These qualitative differences account for the separation of most of the Iberian prehistoric groups from the Central European Neolithic and Early Bronze Age populations along the second component of the PCA, shown on Fig.

We also tested whether this continuity could be the result of data merging across bigger geographic distance. A more detailed analysis of H subhaplogroups focuses on 17 SNPs in the coding region of the human mitogenome. The largest proportion of the H individuals belongs to the subhaplogroup H1 H3 is detected in Chalcolithic individuals from central, southeast and southwest Iberia.

H1 is observed in each period and region, but more frequently in the Chalcolithic and Early Bronze Age than in the Neolithic. The comparative ancient H data from the European prehistory is too sparse for in-depth statistical analyses.

However it becomes apparent from our results that the H diversity in prehistoric Iberia is different from the H diversity of Central Europe 55 , 58 , 67 , 68 , and more similar to the Neolithic populations in France 69 , Overall, we do not observe direct links between the Central European Late Neolithic chronologically comparable to the Chalcolithic in Iberia and the Iberian entire and late CHA groups at the haplogroup level, and thus cannot confirm a Late Chalcolithic expansion toward Central Europe as suggested by haplogroup H mitogenome data The haplotype diversity among the regional groups is the lowest in the SWI Neolithic-Chalcolithic group in Iberia, and that along with the detected common basal J and K haplotypes could cause potential bias in lineage sharing results presented below.

The haplotype sharing SHA between the larger chronological groups detects no hunter-gatherer contribution in the EN group of Iberia, and a low level of hunter-gatherer maternal lineages in the MLN periods 6. During the MLN, several new hunter-gatherer-type U5b lineages appear. In the Chalcolithic period, we also observe a strong ancestral EN contribution The Iberian early farming groups share generally high number of lineages The Central European Early Neolithic has the highest number of shared haplotypes comparing to the northeast Iberian groups, but the sharing is also high with southeast Iberian dataset.

In contrast, lineage sharing with Holocene period hunter-gatherers is the highest in the mostly Chalcolithic southeast Iberian group 8. Concerning temporal succession in NEI region, the proportion of hunter-gatherer lineages increases from 0 to 4.

Continuity between Neolithic and Chalcolithic groups is reflected by the amount of shared lineages between the two periods in CI Genetic distances F st are generally low among the Iberian prehistoric groups, and none of them are significant Supplementary Table S S4 , large-scale relationships are displayed.

Here, the NEI populations are within the cluster of the other Iberian groups, whereas the Central European and French datasets form a cluster that is slightly offset. Interestingly, MDS shows that some groups e. The clustering of the groups on MDS plot was therefore tested by the analysis of molecular variance. The best-supported groupings i. Genetic distances to modern-day populations are generally low, and restricted to certain region s of Europe Supplementary Fig. S5 , Supplementary Table S Italian, French but also Belorussian.

Recent genomic studies have highlighted the similarity of early farmers to modern South Europeans 71 , especially to modern day Sardinians 47 , This theory is based on coalescence date estimates for some mtDNA lineages H1, H3, V, U5b1 , which predate the Neolithic expansion, and also based on the high abundances of these lineages in Iberia today 75 , 76 , 77 , 78 , However, the Franco-Cantabrian glacial refuge theory has not yet been confirmed by ancient mtDNA or genomic analyses, and more Iberian Pleistocene and especially Holocene samples need to be investigated at full mitogenomic resolution.

The indigenous Iberian late Upper Paleolithic and Mesolithic populations were represented by whole mitogenome haplotypes assigned to haplogroup U5b from Northern Spain 43 , The Iberian Peninsula is characterized by diverse landscapes with distinct economic potentials.

During the initial phase of Neolithisation, first farmers probably arrived in northeast Iberia primarily along the Mediterranean route and from there spread along the coastline and rivers e. Our genetic data support a substantial influx of Neolithic immigrants to northeastern Spain, where farmer lineages are most abundant and diverse, and on to other regions, like the SEI and CI. These genetic results match the archaeological data on the mode of Neolithisation described for these areas, so that the fully established Neolithic Iberian communities had distinct hunter-gatherer components 16 , 81 , Geography appears to have been a decisive factor in the advance of Neolithic lifeways.

Prehistoric central Iberia southern and northern Meseta, Ambrona Valley , however, was never an isolated region surrounded by mountains, but rather an important hub for innovations and impulses coming into Iberia On a genetic level, Neolithic central Iberia showed corresponding mitogenomic connections to northeast and southeast Iberian regions Supplementary Table S7.

The Pyrenees are also an area of special interest with regard to the route of immigration of the first farmers. This study includes ten samples from the cave site of Els Trocs, dated to the Early and Middle Neolithic Due to the considerable temporal differences between the occupations, the respective sample sets can be considered as genetically independent.

The 14 C data of the six individuals from the earliest phase cluster closely Supplementary Information. Genome-wide analyses of five of these EN individuals revealed the group to be early Neolithic immigrants 47 , 49 , Therefore, we suspect the presence of a still isolated EN group at Els Trocs.

The isolated geographic position in the Pyrenees suggests immigration of this community from north of the Pyrenees or even from Central Europe via the Rhone Valley rather than from the west along the Mediterranean coastline. This alternative gateway to Spain has also been proposed based on archaeobotanical evidence 17 , The Iberian Middle Neolithic genomic data are still too scarce concentrated on a single site La Mina to draw general conclusions about admixture with late hunter-gatherers La Mina people had According to the data currently available, the homogeneity of Chalcolithic ancient DNA results suggests that human mobility and genetic mixing had generally increased in Iberia by the Chalcolithic.

Interestingly, we did not find evidence for direct genetic links between Chalcolithic Iberia and contemporaneous Central Europe. Links between the two regions had previously been suggested based in particular on Bell Beaker elements present across a wide geographic range 89 , 90 , 91 , as well as by the maternal genetic analyses of the El Mirador site An evaluation of this unexpected observation will need further palaeogenetic studies.

Around BCE, the emergence of the El Argar group was evidently preceded by a break in Chalcolithic cultural traditions in southeast Iberia. Yet there are no apparent new influences or signals of substantial population change on the mtDNA haplogroup level at this time, so that the observed changes may either be due to an upheaval of existing social structures or an influx of groups that cannot be distinguished from the local population at the present level of genetic resolution, e.

Unraveling these apparently contradictory data will certainly require further in-depth analyses both on the archaeological and the archaeogenetic level. The present study, based on new and published mtDNA data of prehistoric Iberian individuals suggests a more complex mode of interaction between local hunter-gatherers and incoming early farmers during the Early and Middle Neolithic of the Iberian Peninsula, as compared to Central Europe.

A characteristic of Iberian population dynamics is the proportion of autochthonous hunter-gatherer haplogroups, which increased in relation to the distance to the Mediterranean coast. In contrast, the early farmers in Central Europe showed comparatively little admixture of contemporaneous hunter-gatherer groups. Already during the first centuries of Neolithic transition in Iberia, we observe a mix of female DNA lineages of different origins. Earlier hunter-gatherer haplogroups were found together with a variety of new lineages, which ultimately derive from Near Eastern farming groups.

On the other hand, some early Neolithic sites in northeast Iberia, especially the early group from the cave site of Els Trocs in the central Pyrenees, seem to exhibit affinities to Central European LBK communities.

The diversity of female linages in the Iberian communities continued even during the Chalcolithic, when populations became more homogenous, indicating higher mobility and admixture across different geographic regions. Even though the sample size available for Early Bronze Age populations is still limited, especially with regards to the El Argar group, we observe no significant changes to the mitochondrial DNA pool until the end of our time transect BCE.

The expansion of groups from the eastern steppe 47 , 92 , which profoundly impacted Late Neolithic and EBA groups of Central and North Europe, cannot yet be seen in the contemporaneous population substrate of the Iberian Peninsula at the present level of genetic resolution. This highlights the distinct character of the Neolithic transition both in the Iberian Peninsula and elsewhere and emphasizes the need for further in-depth archaeogenetic studies for reconstructing the close reciprocal relationship of genetic and cultural processes on the population level.

The studied sites are distributed across the Iberian Peninsula, with an emphasis on the archaeologically relevant regions on the Mediterranean coasts, in central and northern Spain and in southern Portugal Fig. We targeted representative sites from the Mesolithic, Neolithic, Chalcolithic and the Early Bronze Age to cover 4, years of prehistory on the Iberian Peninsula. Sample specific context information were supplied by our colleagues and project partners in Portugal and Spain or collected from previously published papers.

All archaeological sites, relevant radiocarbon dates, and individuals incorporated in the present study are listed in Supplementary Tables S1 - 2. Altogether, individuals from 57 archaeological sites in Spain, Portugal and Morocco were sampled and analyzed for this study. Whenever possible we preferred samples from recent excavations over those that had been held at museum collections for prolonged periods of time.

Here we also report additional samples from the sites of La Mina and Els Trocs. The chronological classification of the burials was based on the archaeological data. In order to avoid possible problems due to terminological inconsistencies, we used temporal from Mesolithic to Early Bronze Age; based on relative chronology and absolute dating and geographic groupings to assess the population mitogenetic data from the Iberian Peninsula. We further distinguished groups based on contextual archaeological evidence, e.

We targeted collecting two to three teeth from each skeleton and sampled bones only when teeth were not available. Samples were then ground to fine powder using a mixer mill Retsch. The milling process was controlled by hydroxylapatite blank controls, which were treated as samples in the subsequent DNA extraction and amplification steps.

Series of samples were tested from each archaeological site, and the amplification success defined any further ancient DNA work. In general, A and B samples from different teeth or bone fragments per individual were analyzed independently. In the sole case is the site of Cova del Cantal, A and B samples, i. Of the total 48 extraction batches, 25 were performed using the phenol-chloroform DNA extraction method 55 and 23 using the silica based DNA extraction protocol We combined different sets of primer pairs for the amplification of the HVS-I of the mitochondrial genome.

Well-preserved samples were amplified in two segments nucleotide position np — , average preserved samples in four fragments np — and samples with very fragmented DNA content were amplified with a combination of six overlapping primer pairs.

PCR, amplicon purification and sequencing protocols were the same as reported in Brandt et al. Note that recent mitochondrial phylogeny updates have complicated the interpretation of typing results: the H1 defining variant GA is now also reported for subhaplogroups H30b, H65a, H79a, Ha Phylotree Build Amplification success was checked via gel electrophoresis.

Individuals with consistent HVS-I profiles from both samples were amplified over the whole HVS-I at least three times from at least two independent extracts, producing between 6—18 independent and overlapping amplicons depending on the primer pairs used. PCR products that showed ambiguous nucleotide positions were cloned to monitor possible background contaminations and DNA damage.

To monitor contamination, sample mix up and other potential errors we used the following authentication strategy: 1. All HVS-I haplotypes were replicated from at least two independent DNA extracts obtained from two skeletal elements from nearly all sampled individuals. Due to the applied primer-systems, amplification of the HVS-I enabled the identification of contiguous sequences with 6—10 SNP calls in the overlapping regions.

The final haplogroup call was based on several independent amplifications targeting different loci of the mitochondrial genome HVS-I and coding region SNPs. All haplotypes could be placed in the mitochondrial phylogeny www. In all cases where ancient haplotypes differed from the current representative haplotypes on the mtDNA phylotree by private mutations, the latter were double-checked by several PCRs and by sequencing in combination with other SNPs.

Altogether 51 of the 2. No contamination could be detected in any of the multiplex PCRs. Of the 51 contaminated blanks, 0. All contaminated blanks were sequenced and compared to haplotypes of samples analyzed along with these PCR extraction blanks.

Fourteen individuals that were amplified from two extraction events had to be excluded from further analyses due to signs of potential contamination. The HVS-I region was sequenced from all relevant co-workers archaeologists, anthropologists and geneticists and compared with samples and blanks Supplementary Table S None of the samples matched with the haplotype of the main processor of the samples C.

However, we find no signs of systematic contamination by these workers and consider it unlikely that samples that share these haplotypes were affected in various independent experiments whilst parallel samples were not. Ten samples were captured for whole mitogenomes 47 , and these results corresponds to the PCR and clone based sequence results Supplementary Table S The dataset from Cova de Montanissel was only used in haplogroup based tests, due to insufficiently reproduced HVS-I results Where the chronology could not be defined precisely, we classified the samples into Neolithic Neo and Chalcolithic CHA periods.

We used principal component analysis PCA to visualize the relationships between relative haplogroup compositions of the Iberian and other prehistoric datasets. PCA has the benefit of reducing the complexity of the entire dataset while maintaining an accurate representation of its variability.

Hierarchical clustering was used to construct clusters of the predefined prehistoric datasets, based on similarities distances between their haplogroup compositions.

This analysis helps interpreting the PCA plots in form of dendrograms. Clustering of haplogroup frequencies was performed using Ward type algorithm and Euclidean distance measurement method. The result was visualized as a dendogram with the pvclust package in R. Cluster significance was evaluated by 10, bootstrap replicates. Significance of each cluster was given as an AU Approximately Unbiased p-value, in percentage.

The resulting p-value thus described the probability of obtaining the observed haplogroup compositions if both groups were part of the same metapopulation. We used all haplogroup results per group in this analyses and tested a series of pairwise comparisons of Iberian chronological and geographic groups, using fisher.

The test of population continuity explores whether genetic differences between two populations from two or more consecutive time periods can be adequately explained by genetic drift alone or whether a discontinuity between the two populations has to be assumed The tests were performed for pairs of chronological groups in specific regions central Iberia, northeast Iberia.

Further parameters of the tests are presented in Supplementary Table S8. We also characterized the Iberian populations with standard diversity indices that were calculated in DnaSP Shared haplotype analysis examines the occurrence of identical lineages in different populations. Levelplot of the percentage of shared lineages was visualized in R version 3.



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