Jump to content

Genetic history of Italy

From Wikipedia, the free encyclopedia

Part of a series on the
History of Italy
Old map of Italian peninsula
Early
Ancient Rome
Romano-Barbarian Kingdoms
Odoacer's 476–493
Ostrogothic 493–553
Vandal 435–534
Lombard 568–774
Frankish (Carolingian Empire) 774–962
Germanic (Holy Roman Empire) 962–1801
Medieval
Early modern
Modern
By topic

Timeline

flag Italy portal
Principal Component Analysis of the Italian population[1]

The genetic history of Italy includes information around the formation, ethnogenesis, and other DNA-specific information about the inhabitants of Italy. Modern Italians mainly descend from the ancient peoples of Italy, including Indo-European speakers (Romans and other Latins, Falisci, Picentes, Umbrians, Samnites, Oscans, Sicels, Elymians, Messapians and Adriatic Veneti, as well as Magno-Greeks, Cisalpine Gauls and Illyric Iapygians) and pre-Indo-European speakers (Etruscans, Ligures, Rhaetians Camunni, Sicani, Nuragic peoples), as well as settlers from Phoenicia and Carthage in Sardinia and in the westernmost part of Sicily. Other groups migrated into Italy as a result of the Roman Empire, when the Italian peninsula attracted people from the various regions of the empire (North Africa, West Asia and the rest of Europe),[2] and during the Middle Ages with the arrival of Ostrogoths, Longobards, Franks, Saracens, and Normans among others. Based on DNA analysis, there is evidence of regional genetic substructure and continuity within modern Italy dating back to antiquity.[3][4][5][6]

In their admixture ratios, Italians are similar to other Southern Europeans, and that is being of primarily Neolithic Early European Farmer ancestry, along with smaller, but still significant, amounts of Mesolithic Western Hunter-Gatherer, Bronze Age Steppe pastoralist (Indo-European speakers) and Chalcolithic or Bronze Age Iranian/Caucasus-related ancestry.[4][7][8][9][10] According to multiple genome-wide studies Southern Italians are closest to modern Greeks,[11][12][13][14][15][16][17][18][19][10] while Northern Italians are closest to the Spaniards, the Portuguese and to a lesser extent people from Southern France.[11][18][20][21][22][17][23][24][25][26][19][27] There is also Bronze/Iron Age Anatolian admixture in Italy, with a much lower incidence in Northern Italy compared with Central Italy and Southern Italy.[24][8] Marginal levels (0.1-4%) of North African admixture is also found in Southern Italy with the highest incidence being in the islands of Sicily and Sardinia.[24][8][4]

Overview

[edit]

Latin samples from Latium in the Iron Age and early Roman Republican period were generally found to genetically cluster closest to modern Northern and Central Italians (four out of six were closest to Northern and Central Italians, while the other two were closest to Southern Italians).[28] DNA analysis demonstrates that ancient Greek colonization had a significant lasting effect on the local genetic landscape of Southern Italy and Sicily (Magna Graecia), with modern people from that region having significant Greek admixture.[29][14] Overall, the genetic differentiation between the Latins, Etruscans and the preceding proto-Villanovan population of Italy was found to be insignificant.[30] In 2019, aDNA analysis of Roman fossils detected substantial genetic ancestry shift towards central and northern European ancestry in the inhabitants of the city of Rome in late antiquity and the medieval era. The authors tentatively link the origin of this ancestry with Visigoths and Lombards. Previously, most citizens in the imperial era clustered with central and east Mediterranean peoples, such as central and south Italians, Greeks, Cypriots and Maltese, and to some extent, Levantine and Near Eastern peoples. This was caused by direct immigration and contact with Greek, Phoenician and Punic diasporas.[2][31] A 2020 analysis of maternal haplogroups from ancient and modern samples indicates a substantial genetic similarity and continuity between the modern inhabitants of Umbria in central Italy and ancient inhabitants of the region belonging to the Italic-speaking Umbrian culture.[6]

Multiple DNA studies confirmed that genetic variation in Italy is clinal, going from the Eastern to the Western Mediterranean. The Sardinians are the exception as genetic outliers in Italy and indeed in Europe, resulting from their predominantly Neolithic, Pre-Indo-European and non-Italic Nuragic ancestry.[32][7][8][9] Reflecting the history of Europe and the broader Mediterranean basin, the Italian populations have been found to be made up mostly of the same ancestral components, albeit in different proportions, from the Mesolithic, Neolithic and Bronze Age settlements of Europe.[24][21][33]

The genetic gap between northern and southern Italians is filled by an intermediate Central Italian cluster, creating a continuous cline of variation that mirrors geography.[25] The only exceptions are some minority populations (mostly Slovene minorities from the region of Friuli-Venezia Giulia) who cluster with the Slavic-speaking Central Europeans in Slovenia,[34] as well as the Sardinians, who are clearly differentiated from the populations of both mainland Italy and Sicily.[21][4] A study on some linguistic and isolated communities residing in Italy revealed that their genetic diversity at short (0–200 km) and intermediate distances (700–800 km) was greater than that observed throughout the entire European continent.[5]

The genetic distance between Northern and Southern Italians, although large for a single European nationality, is similar to that between the Northern and the Southern Germans.[35] Northern and Southern Italians began to diverge as early as the Late Glacial, and appear to encapsulate at a smaller scale the cline of genetic diversity observable across Europe.[36]

Prehistoric and historical populations of Italy

[edit]
Timeline summarizing the main demographic events in ancient Italy, from the Paleolithic to the Middle Ages.
Ethnic groups of Italy (as defined by today's borders) in the 4th century BC.

Modern humans appeared during the Upper Paleolithic. Specimens of Aurignacian age were discovered in the cave of Fumane and date back about 34,000 years. During the Magdalenian period, the first humans from the Pyrenees populated Sardinia.[37]

During the Neolithic period, farming was introduced by people from the east and the first villages were built. Weapons became more sophisticated and the first objects in clay were produced. In the late Neolithic era the use of copper spread, and villages were built over piles near lakes. In Sardinia, Sicily and a part of Mainland Italy the Beaker culture spread from Western and Central Europe. Sicily also suffered the influences of the Aegean in the Mycenaean period.

During the Late Bronze Age the Urnfield Proto-Villanovan culture appeared in Central and Northern Italy. It is characterized by the rite of cremation of dead bodies, which originated from Central Europe. The use of iron began to spread.[38] In Sardinia, the Nuragic civilization flourished.

At the dawn of the Iron Age much of Italy was inhabited by Italic tribes such as the Latins, Sabines, Samnites, and Umbrians. The Northwest and Alpine territories were populated primarily by pre-Indo European speakers such as the Etruscans, Ligurians, Camunni and Raetians; while Iapygian tribes, possibly of Illyrian origin, populated Apulia.

From the 8th century BC, Greek colonists settled on the southern Italian coast and founded cities, forming what would be later called Magna Graecia. Around the same time, Phoenician colonists settled on the western side of Sicily. During the same period the Etruscan civilization developed on the coast of Southern Tuscany and Northern Latium. In the 4th century BC, Gauls settled in Northern Italy and in parts of Central Italy.[39] With the fall of the Western Roman Empire, different populations of Germanic origin invaded Italy, the most significant being the Lombards,[40] followed five centuries later by the Normans in Sicily.

Daunians

[edit]
This section is an excerpt from Daunians § Genetics.[edit]

A genetic study published in 2022 examined DNA extracted from three necropoleis: Ordona, Salapia and San Giovanni Rotondo, in the region of Apulia in southern Italy, which during the Iron Age have been linked to the Daunian region. Most samples from Ordona and Salapia date to the Daunian period and some samples from San Giovanni Rotondo date more broadly to the Iron Age. Paternal haplogroups of seven Iron Age samples were identified. Two paternal lineages of the Iron Age samples belong to J-M241, one of them could be further processed as J-L283+. Two Iron Age samples belonged to R-M269, one further designated as Z2103+ and one to I-M223.[41]

Iron Age Daunians showed the highest autosomal affinity with Early Iron Age Illyrian populations from Croatia and populations which were formed in Italy in the Roman Republican era, which both can be broadly included in a pan-Mediterranean genetic continuum (stretching from Crete to Republican Rome and the Iberian Peninsula). Links to Minoans/Crete and Iron Age Greeks/Arkadia are less likely. A parsimonious explanation of the Daunian's origin favors a genetic continuity between the Daunians and the population that inhabited the area prior to the historical period that was analyzed, although additional influences from Croatia (ancient Illyria) cannot be excluded, as described by the material remains and the available historical sources.[42]

Etruscans

[edit]
This section is an excerpt from Etruscan origins § Genetic evidence.[edit]
Etruscan votive heads, IV-II century BC

There have been numerous biological studies on the Etruscan origins, the oldest of which dates back to the 1950s when research was still based on blood tests of modern samples, and DNA analysis (including the analysis of ancient samples) was not yet possible.[43][44][45] It is only in very recent years, starting in 2019, with the development of archaeogenetics, that comprehensive studies containing the whole genome sequencing (WGS) of Etruscan samples have been published, including autosomal DNA and Y-DNA, autosomal DNA being the "most valuable to understand what really happened in an individual's history", as stated by geneticist David Reich, whereas previously studies were based only on mitochondrial DNA analysis, which contains less and limited information.[46] The direct testing of ancient Etruscan DNA supports a deep, local origin, while the testing of modern samples as a proxy for Etruscans has proven to be rather inconclusive and inconsistent.[47][48]

A 2019 genetic study published in the journal Science analyzed the autosomal DNA of 11 Iron Age samples from the areas around Rome, including for the first time the whole genome sequencing (WGS) of some samples from Etruscan tombs, and concluded that Etruscans (900-600 BC) and the Latins (900-200 BC) from Latium vetus were genetically similar, and Etruscans also had Steppe-related ancestry despite speaking a pre-Indo-European language.[49] A 2021 genetic study published in the journal Science Advances analyzed the autosomal DNA of 48 Iron Age individuals from Tuscany and Lazio and confirmed that the Etruscan individuals displayed the ancestral component Steppe in the same percentages as found in the previously analyzed Iron Age Latins, and that the Etruscans' DNA completely lacks a signal of recent admixture with Anatolia or the Eastern Mediterranean, concluding that the Etruscans were autochthonous and they had a genetic profile similar to their Latin neighbors. Both Etruscans and Latins joined firmly the European cluster, 75% of the Etruscan male individuals were found to belong to haplogroup R1b-M269 and its subclades, especially R1b-P312 and its derivative R1b-L2 whose direct ancestor is R1b-U152, while the most common mitochondrial DNA haplogroup among the Etruscans was H.[50] Iron Age Etruscans from central Italy could be modelled as deriving 50% of their ancestry from Central European Bell Beakers, represented by Germany Bell Beaker, with around 25-30% Steppe-related ancestry, and the rest of their ancestry from a local Chalcolithic population. The conclusions of these studies have been confirmed by later ones.[51]

In the collective volume Etruscology published in 2017, British archeologist Phil Perkins provides an analysis of the state of DNA studies and writes that "none of the DNA studies to date conclusively prove that Etruscans were an intrusive population in Italy that originated in the Eastern Mediterranean or Anatolia" and "there are indications that the evidence of DNA can support the theory that Etruscan people are autochthonous in central Italy".[48]

In his book A Short History of Humanity published in 2021, German geneticist Johannes Krause, co-director of the Max Planck Institute for Evolutionary Anthropology in Jena, concludes that it is likely that the Etruscan language (as well as Basque, Paleo-Sardinian and Minoan) "developed on the continent in the course of the Neolithic Revolution".[52]

Latins

[edit]
Main article: Latins (Italic tribe)

A genetic study published in Science in November 2019 examined the remains of six Latin males buried near Rome between 900 BC and 200 BC. They carried the paternal haplogroups R-M269, T-L208, R-P311, R-PF7589 and R-P312 (two samples), and the maternal haplogroups H1aj1a, T2c1f, H2a, U4a1a, H11a and H10.[53] These examined individuals were distinguished from preceding populations of Italy by the presence of 30% steppe ancestry.[54] Two out of six individuals from Latin burials were found have a mixture of local Iron Age ancestry and ancestry from an Eastern Mediterranean population. Among modern populations, four out of six were closest to Northern and Central Italians, and then Spaniards, while the other two were closest to Southern Italians.[28] Overall, the genetic differentiation between the Latins, Etruscans and the preceding proto-Villanovan population of Italy was found to be insignificant.[30]

Nuragic civilization

[edit]
This section is an excerpt from Nuragic civilization § Paleogenetics.[edit]

A genetic study published in Nature Communications in February 2020 examined the remains of 17 individuals identified with Nuragic civilization. The samples of Y-DNA extracted belonged to haplogroup I2a1b1 (2 samples), R1b1b2a, G2a2b2b1a1, R1b1b (4 samples), J2b2a1 (3 samples) and G2a2b2b1a1a, while the samples of mtDNA extracted belonged to various types of haplogroup T, V, H, J, K and U.[55] The study found strong evidence of genetic continuity between Nuragic civilization and earlier Neolithic inhabitants of Sardinia, who were genetically similar to Neolithic peoples of Iberia and southern France.[56] They were determined to be of about 80% Early European Farmer (EEF) ancestry and 20% Western Hunter-Gatherer (WHG) ancestry.[57] They were predicted to be largely descended from peoples of the Neolithic Cardial Ware culture, which spread throughout the western Mediterranean in Southern Europe c. 5500 BC.[58] The Nuragic people were differentiated from many other Bronze Age peoples of Europe by the near absence of steppe-related ancestry.[56][59]

A 2021 study by Villalba-Mouco et al. has identified a possible gene flow originating from the Italian peninsula starting from the Chalcolithic. In prehistoric Sardinia, the component associated with Iranian farmers, or Caucasus-related ancestry, present in Mainland Italy since the Neolithic (together with the EEF and WHG components), gradually increases from 0% in the Early Chalcolithic to about 5.8% in the Bronze Age.[60] The absence of the component linked to the Magdalenians would instead exclude contributions from the Chalcolithic of the south of the Iberian peninsula.[60] According to a 2022 study by Manjusha Chintalapati et al., "In Sardinia, a majority of the Bronze Age samples do not have Steppe pastoralist-related ancestry. In a few individuals, we found evidence for steppe ancestry", which would arrive in ~2600 BC.[61]

Genetic data appears to support the hypothesis of a patrilocal society. In various sites, more genetic diversity was found in the mitochondrial DNA, suggesting that women may have moved from community to community more than the men.[62]

Picentes

[edit]
This section is an excerpt from Picentes § Genetics.[edit]

A 2017 analysis of maternal haplogroups from ancient and modern samples indicated a substantial genetic similarity among the modern inhabitants of Central Italy and the area's ancient pre-Roman inhabitants of settlement of Novilara in the province of Pesaro, and evidence of substantial genetic continuity in the region from pre-Roman times to the present with regard to mitochondrial DNA.[63]

In a 2024 study of Iron Age ancient samples, the two main Y haplogroups in Picene were R1-M269/L23 (58% of the total) and J2-M172/M12 (25% of the total), which respectively indicate the direct relationship with Central Europe and the Balkan peninsula. Particularly, the R1-M269/L23 haplogroup was related to the Yamnaya ancestry and was present at high frequency in Central European populations since the Bronze Age onwards.[64]

Lombards

[edit]
This section is an excerpt from Lombards § Genetics.[edit]

A genetic study published in Nature Communications in September 2018 found strong genetic similarities between Lombards of Italy and earlier Lombards of Central Europe. Lombard males were primarily carriers of subclades of haplogroup R1b and I2a2a1, both of which are common among Germanic peoples. Lombard males were found to be more genetically homogeneous than Lombard females. The evidence suggested that the Lombards originated in Central/Northern Europe, and were a patriarchal people who settled Central Europe and then later Italy through a migration from the north.[65]

A genetic study published in Science Advances in September 2018 examined the remains of a Lombard male buried at an Alemannic graveyard. He was found to be a carrier of the paternal haplogroup R1b1a2a1a1c2b2b and the maternal haplogroup H65a. The graveyard also included the remains of a Frankish and a Byzantine male, both of whom were also carriers of subclades of the paternal haplogroup R1b1a2a1a1. The Lombard, Frankish and Byzantine males were all found to be closely related, and displayed close genetic links to Northern Europe, particularly Lithuania and Iceland.[66]

A genetic study published in the European Journal of Human Genetics in January 2019 examined the mtDNA of a large number of early-medieval Lombard remains from Central Europe and Italy. These individuals were found to be closely related and displayed strong genetic links to Central Europe. The evidence suggested that the Lombard settlement of Italy was the result of a migration from the north involving both males and females.[67]

A 2024 paper found that the Lombards of Italy were best modelled by a Jutlandic Iron Age source, consistent with an origin in Jutland or northern Germany.[68]

Y-DNA genetic diversity

[edit]
Main article: Y-DNA haplogroups in populations of Europe
Distribution of the R1b haplogroup in Europe

Many Italians, especially in Northern Italy and Central Italy, belong to Haplogroup R1b, common in Western and Central Europe. The highest frequency of R1b is found in Garfagnana (76.2%) in Tuscany and in the Bergamo Valleys (80.8%) in Lombardy.[69][70] This percentage lowers in the south of Italy in Calabria (33.2%).[69] On the other hand 39% of the Sardinians belong to Mesolithic European haplogroup I2a1a.[71][72]

A study from the Università Cattolica del Sacro Cuore found that while Greek colonization left little significant genetic contribution, data analysis sampling 12 sites in the Italian peninsula supported a male demic diffusion model and Neolithic admixture with Mesolithic inhabitants.[73] The results supported a distribution of genetic variation along a north–south axis and supported demic diffusion. South Italian samples clustered with southeast and south-central European samples, and northern groups with West Europe.[73][74]

A 2004 study by Semino et al. showed that Italians from the north-central regions had around 26.9% J2; the Apulians, Calabrians and Sicilians had 29.1%, 21.5% and 16.7% J2 respectively; the Sardinians had 9.7% J2.[75]

A 2018 genetic study, focusing on the Y-chromosome and haplogroups lineages, their diversity and their distribution by taking some 817 representative subjects, gives credit to the traditional northern-southern division in population, by concluding that due to Neolithic migrations southern Italians "show a higher similarity with Middle Eastern and Southern Balkan populations than northern ones; conversely, northern samples are genetically closer to North-West Europe and Northern Balkan groups". The position of Volterra in central Tuscany keeps the debate about the origins of Etruscans open, although the numbers are strongly in favor of the autochthonous thesis: the low presence of J2a-M67* (2.7%) suggests contacts by sea with Anatolian people; the presence of Central European lineage G2a-L497 (7.1%) at considerable frequency would rather support a Central European origin of the Etruscans; and finally, the high incidence of European R1b lineages (R1b 50% approx., R1b-U152 24.5%) — especially of haplogroup R1b-U152 — could suggest an autochthonous origin due to a process of formation of the Etruscan civilisation from the preceding Villanovan culture, following the theories of Dionysius of Halicarnassus,[69] as already supported by archaeology, anthropology and linguistics.[76][77][78][79] In 2019, in a Stanford study published in Science, two ancient samples from the Neolithic settlement of Ripabianca di Monterado in province of Ancona, in the Marche region of Italy, were found to be Y-DNA J-L26 and J-M304.[2] Therefore, Y-DNA J2a-M67, downstream to Y-DNA J-L26 and J-M304, is most likely in Italy since the Neolithic and can't be the proof of recent contacts with Anatolia.

Y-DNA introduced by historical immigration

[edit]

In two villages in Lazio and Abruzzo (Cappadocia and Vallepietra), I1 is the most common Y-DNA, recorded at levels 35% and 28%.[80] In Sicily, further migrations from the Vandals and Saracens have only slightly affected the ethnic composition of the Sicilian people. However, specifically Greek genetic legacy is estimated at 37% in Sicily.[29]

The Norman conquest of southern Italy caused the Norman Kingdom of Sicily to be created in 1130, with Palermo as capital, 70 years after the initial Norman invasion and 40 after the conquest of the last town, Noto in 1091, and would last until 1198. Nowadays it is in central and western Sicily, that Norman Y-DNA is common, with 15% to 20% of the lineages belonging to haplogroup I, this percentage drops to 8% in the eastern part of the island. The North African male contribution to Sicily was estimated between 0% and 7.5%.[81][14][82] Overall, the estimated Southern Balkan and Western European paternal contributions in Sicily are about 63% and 26% respectively.[82]

A 2015 genetic study of six small mountain villages in eastern Lazio and one mountain community in nearby western Abruzzo found some genetic similarities between these communities and Near Eastern populations, mainly in the male genetic pool. The Y haplogroup Q, common in Western Asia and Central Asia, was also found among this sample population, suggesting that in the past could have hosted a settlement from Anatolia.[80] Also, it is about 0.6% in continental Italy, but it rises to 2.5% (6/236) in Sicily, where it reaches 16.7% (3/18) in Mazara del Vallo region, followed by 7.1% (2/28) in Ragusa, 3.6% in Sciacca,[29] and 3.7% in Belvedere Marittimo.[83]

Genetic composition of Italian mtDNA

[edit]

In Italy as elsewhere in Europe the majority of mtDNA lineages belong to the haplogroup H. Several independent studies conclude that haplogroup H probably evolved in West Asia c. 25,000 years ago. It was carried to Europe by migrations c. 20–25,000 years ago, and spread with population of the southwest of the continent.[84][85] Its arrival was roughly contemporary with the rise of the Gravettian culture. The spread of subclades H1, H3 and the sister haplogroup V reflect a second intra-European expansion from the Franco-Cantabrian region after the last glacial maximum, c. 13,000 years ago.[84][86]

African Haplogroup L lineages are relatively infrequent (less than 1%) throughout Italy with the exception of Latium, Volterra, Basilicata and Sicily where frequencies between 2 and 3% have been found.[87]

A study in 2012 by Brisighelli et al. stated that an analysis of ancestral informative markers "as carried out in the present study indicated that Italy shows a very minor sub-Saharan African component that is, however, slightly higher than non-Mediterranean Europe." Discussing African mtDNAs the study states that these indicate that a significant proportion of these lineages could have arrived in Italy more than 10,000 years ago; therefore, their presence in Italy does not necessarily date to the time of the Roman Empire, the Atlantic slave trade or to modern migration."[33] These mtDNAs by Brisighelli et al. were reported with the given results as "Mitochondrial DNA haplotypes of African origin are mainly represented by haplogroups M1 (0.3%), U6 (0.8%) and L (1.2%) for the 583 samples tested.[33] The haplogroups M1 and U6 can be considered to be of North African origin and could therefore be used to signal the documented African historical input. Haplogroup M1 was observed in only two carriers from Trapani (West Sicily), while U6 was observed only in Lucera, South Apulia, and another at the tip of the Peninsula (Calabria).[33]

A 2013 study by Alessio Boattini et al. found 0 of African L haplogroup in the whole Italy out of 865 samples. The percentages for Berber M1 and U6 haplogroups were 0.46% and 0.35%, respectively.[88]

A 2014 study by Stefania Sarno et al. found 0 of African L and M1 haplogroups in mainland Southern Italy out of 115 samples. Only two Berber U6 out of 115 samples were found, one from Lecce and one from Cosenza.[82]

A close genetic similarity between Ashkenazim and Italians has been noted in genetic studies, possibly due to the fact that Ashkenazi Jews have a significant European admixture (30–60%), much of it Southern European, a lot of which came from Italy when Jewish diaspora males of Middle Eastern origin migrated to Rome and found wives among local women who then converted to Judaism.[89][90][91][92][93][22][94] More specifically, Ashkenazi Jews could be modeled as being 50% Levantine and 50% European, with an estimated mean South European admixture of 37.5%. Most of it (30.5%) seems to derive from an Italian source.[95][96]

A 2010 study of Jewish genealogy found that with respect to non-Jewish European groups, the populations which are most closely related to Ashkenazi Jews are modern-day Italians followed by the French and Sardinians.[97][98]

Recent studies have shown that Italy played an important role in the recovery of "Western Europe" at the end of the Last glacial period. The study which was focused on the mitochondrial U5b3 haplogroup discovered that this female lineage had in fact originated in Italy and around 10,000 years ago it expanded from the Peninsula towards Provence and the Balkans. In Provence, probably between 9,000 and 7,000 years ago, it gave rise to the haplogroup subclade U5b3a1. This subclade U5b3a1 later came from Provence to the island of Sardinia by way of obsidian merchants, because it is estimated that 80% of the obsidian which is found in France comes from Monte Arci in Sardinia, reflecting the close relationship which once existed between these two regions. Still about 4% of the female population of Sardinia belongs to this haplotype.[99]

A mtDNA study, published in 2018 in the journal American Journal of Physical Anthropology, compared both ancient and modern samples from Tuscany, from the Prehistory, Etruscan age, Roman age, Renaissance, and Present-day, and concluded that the Etruscans appear as a local population, intermediate between the prehistoric and the other samples, placing in the temporal network between the Eneolithic Age and the Roman Age.[100]

A 2020 analysis of maternal haplogroups from ancient and modern samples in the central Italian region of Umbria finds a substantial genetic similarity among modern Umbrians and the area's pre-Roman inhabitants, and evidence of substantial genetic continuity in the region from pre-Roman times to the present. Both modern and ancient Umbrians were found to have high rates of mtDNA haplogroups U4 and U5a, and an overrepresentation of J (at roughly 30%). The study also found that, "local genetic continuities are further attested to by six terminal branches (H1e1, J1c3, J2b1, U2e2a, U8b1b1 and K1a4a)" also shared by ancient and modern Umbrians.[6]

Autosomal

[edit]
Admixture plots of modern West Eurasian populations based on seven components:
     South/West European      North/East European      Caucasus
     West Asian      South Asian      East Asian      North African/Sub-Saharan African[101]
The European genetic structure (based on 273,464 SNPs). Three levels of structure as revealed by PC analysis are shown: A) inter-continental; B) intra-continental; and C) inside a single country (Estonia), where median values of the PC1&2 are shown. D) European map illustrating the origin of sample and population size. CEU – Utah residents with ancestry from Northern and Western Europe, CHB – Han Chinese from Beijing, JPT – Japanese from Tokyo, and YRI – Yoruba from Ibadan, Nigeria.[35]

Wade et al. (2008) determined that Italy is one of the last two remaining genetic islands in Europe, the other being Finland. This is due in part to the presence of the Alpine mountain chain which, over the centuries, has prevented large migration flows.[102]

Recent genome-wide studies have been able to detect and quantify admixture like never before. Li et al. (2008), using more than 600,000 autosomal SNPs, identify seven global population clusters, including European, Middle Eastern and Central/South Asian. All the Italian samples belong to Central-Western group with minor influences dating to Neolithic period.[103]

López Herráez et al. (2009) typed the same samples at close to 1 million SNPs and analyzed them in a Western Eurasian context, identifying a number of subclusters. This time, all of the European samples show some minor admixture. Among the Italians, Tuscany still has the most, and Sardinia has a bit too, but so does Lombardy (Bergamo), which is even farther north.[104]

A 2011 study by Moorjani et al. found that many southern Europeans have inherited 1–3% Sub-Saharan ancestry, although the percentages were lower (0.2–2.1%) when reanalyzed with the 'STRUCTURE' statistical model. An average admixture date of around 55 generations/1100 years ago was also calculated, "consistent with North African gene flow at the end of the Roman Empire and subsequent Arab migrations"[105]

A 2012 study by Di Gaetano et al. used 1,014 Italians with wide geographical coverage. It showed that the current population of Sardinia can be clearly differentiated genetically from mainland Italy and Sicily, and that a certain degree of genetic differentiation is detectable within the current Italian peninsula population.

By using the ADMIXTURE software, the authors obtained at K = 4 the lowest cross-validation error. The HapMap CEU individuals showed an average Northern Europe (NE) ancestry of 83%. A similar pattern is observed in French, Northern Italian and Central Italian populations with a NE ancestry of 70%, 56% and 52% respectively. According to the PCA plot, also in the ADMIXTURE analysis there are relatively small differences in ancestry between Northern Italians and Central Italians while Southern Italians showed a lower average admixture NE proportion (44%) than Northern and Central Italy, and a higher Caucasian ancestry of 28%. The Sardinian samples display a pattern of crimson common to the others European populations but at a higher frequency (70%). The average admixture proportions for Northern European ancestry within current Sardinian population is 14.3% with some individuals exhibiting very low Northern European ancestry (less than 5% in 36 individuals on 268 accounting the 13% of the sample).[21]

A 2013 study by Peristera Paschou et al. confirms that the Mediterranean Sea has acted as a strong barrier to gene flow through geographic isolation following initial settlements. Samples from (Northern) Italy, Tuscany, Sicily and Sardinia are closest to other Southern Europeans from Iberia, the Balkans and Greece, who are in turn closest to the Neolithic migrants that spread farming throughout Europe, represented here by the Cappadocian sample from Anatolia. But there hasn't been any significant admixture from the Middle East or North Africa into Italy and the rest of Southern Europe since then.[17]

Ancient DNA analysis reveals that Ötzi the Iceman clusters with modern Southern Europeans and closest to Italians (the orange "Europe S" dots in the plots below), especially those from the island of Sardinia. Other Italians pull away toward Southeastern and Central Europe consistent with geography and some post-Neolithic gene flow from those areas (e.g. Italics, Greeks, Etruscans, Celts), but despite that and centuries of history, they're still very similar to their prehistoric ancestor.[106]

A 2013 study by Botigué et al. 2013 applied an unsupervised clustering algorithm, ADMIXTURE, to estimate allele-based sharing between Africans and Europeans. Regarding Italians, the North African ancestry does not exceed 2% of their genomes. On average, 1% of Jewish ancestry is found in Tuscan HapMap population and Italian Swiss, as well as Greeks and Cypriots. Contrary to past observations, Sub-Saharan ancestry is detected at <1% in Europe, with the exception of the Canary Islands.[82]

Haak et al. (2015) conducted a genome wide study of 94 ancient skeletons from Europe and Russia. The study argues that Bronze Age steppe pastoralists from the Yamna culture spread Indo-European languages in Europe. Autosomic tests indicate that the Yamnaya-people were the result of admixture between two different hunter-gatherer populations: Eastern Hunter-Gatherers from the Russian Steppe and either Caucasus Hunter-Gatherers or Chalcolithic Iranians (who are very similar). Wolfgang Haak estimated a 27% ancestral contribution of the Yamnaya in the DNA of modern Tuscans, a 25% ancestral contribution of the Yamnaya in the DNA of modern Northern Italians from Bergamo, excluding Sardinians (7%), and to a lesser extent Sicilians (12%).[20]

A 2016 study Sazzini et al., confirms the results of previous studies by Di Gaetano et al. (2012) and Fiorito et al. (2015) but has much better geographical coverage of samples, with 737 individuals from 20 locations in 15 different regions being tested. The study also for the first time includes a formal admixture test that models the ancestry of Italians by inferring admixture events using all of the Western Eurasian samples. The results are very interesting in light of the ancient DNA evidence that has come out in the last couple years:

In addition to the pattern described in the main text, the SARD sample seemed to have played a major role as source of admixture for most of the examined populations, especially Italian ones, rather than as recipient of migratory processes. In fact, the most significant f3 scores for trios including SARD indicated peninsular Italians as plausible results of admixture between SARD and populations from Iran, Caucasus and Russia. This scenario could be interpreted as further evidence that Sardinians retain high proportions of a putative ancestral genomic background that was considerably widespread across Europe at least until the Neolithic and that has been subsequently erased or masked in most of present-day European populations.[25]

Sarno et al. (2017) concentrate on the genetic impact brought by the historical migrations around the Mediterranean on Southern Italy and Sicily, and conclude that the "results demonstrate that the genetic variability of present-day Southern Italian populations is characterized by a shared genetic continuity, extending to large portions of central and eastern Mediterranean shores", while showing that "Southern Italy appear more similar to the Greek-speaking islands of the Mediterranean Sea, reaching as far east as Cyprus, than to samples from continental Greece, suggesting a possible ancestral link which might have survived in a less admixed form in the islands", also precises how "besides a predominant Neolithic-like component, our analyses reveal significant impacts of Post-Neolithic Caucasus- and Levantine-related ancestries."[14] A news article associated with the Max Planck Society, reviewing the results, while beginning by stating that "populations along the eastern Mediterranean coast share a genetic heritage that transcends nationality", also points out how this study is interesting on the debates concerning the diffusion of the Indo-European languages family in Europe, as, while showcasing the influence from the Caucasus, there's no genetic marker associated with the Pontic–Caspian steppe, "a very characteristic genetic signal well represented in North-Central and Eastern Europe, which previous studies associated with the introduction of Indo-European languages to the continent."[14]

Raveane et al. (2019) discovered in a genome-wide study on modern-day Italians a contribution of Caucasus Hunter-Gatherers from the third millennium Anatolian Bronze Age, predominantly in Southern Italy. Furthermore, patterns of regional variation showed geographical structure in Southern Italy, Northern Italy, and Sardinia, in line with previous studies. Even more detailed structure was observed between subregional clusters, caused by geography and distance, and historical admixture possibly associated with events at the end of the Roman Empire and during subsequent periods.[4]

Antonio et al. (2019) studied historical populations from various time periods in Latium and Rome. They found that, despite the linguistic differences, the Latins and the Etruscans showed no significant genetic differences. Their autosomal DNA was a mixture in similar proportions of Western Hunter-Gatherers (Mesolithic), Early European Farmers (Neolithic), and Western Steppe Herders (Bronze Age).[2]

A 2022 genome-wide study of more than 700 individuals from the South Mediterranean area (102 from Southern Italy), combined with ancient DNA from neighbouring areas, found high affinities of South-Eastern Italians with modern Eastern Peloponnesians, and a closer affinity of ancient Greek genomes with those from specific regions of South Italy than modern Greek genomes. The study also discovered common genetic sources shared between South Italy and Peloponnese, which can be modeled as a mixture of Anatolian Neolithic and Iranian Chalcolithic ancestries.[108]

A 2023 study examined the Balkan samples from the Roman era, finding surprisingly little ancestry contribution from peoples of Italic descent.[127]

Monnereau et al (2024) analyzed burials at the site of Segesta to investigate the interactions between Muslim and Christian communities during the Middle Ages in Sicily. The biomolecular and Isotopic results suggest the Christians remained genetically distinct from the Muslim community at Segesta while following a substantially similar diet. Based on these results, the authours suggest that the two communities at Segesta could have followed endogamy rules.[128]

Baker et al. (2024) used implement typology to divide the Gravettian technocomplex into nine distinct groups, including two closely related groups in Italy, comparing these with the archaeogenetic evidence,[137] most notably from Allentoft et al. (2022).[165]

A 2024 study analysed the genomic ancestry and social dynamics of Western Hunter-Gatherers, including several from Italy.[177]

Another 2024 study sequenced human-animal co-burials from the Late Iron Age necropolis of the Seminario Vescovile in Verona.[187]

See also

[edit]

References

[edit]
  1. ^ Parolo S, Lisa A, Gentilini D, Di Blasio AM, Barlera S, Nicolis EB, et al. (November 2015). "Characterization of the biological processes shaping the genetic structure of the Italian population". BMC Genetics. 16: 132. doi:10.1186/s12863-015-0293-x (inactive 1 December 2024). PMC 4640365. PMID 26553317.{{cite journal}}: CS1 maint: DOI inactive as of December 2024 (link)
  2. ^ a b c d Antonio ML, Gao Z, Moots HM, Lucci M, Candilio F, Sawyer S, et al. (November 2019). "Ancient Rome: A genetic crossroads of Europe and the Mediterranean". Science. 366 (6466). American Association for the Advancement of Science (published November 8, 2019): 708–714. Bibcode:2019Sci...366..708A. doi:10.1126/science.aay6826. hdl:2318/1715466. PMC 7093155. PMID 31699931. Interestingly, although Iron Age individuals were sampled from both Etruscan (n=3) and Latin (n=6) contexts, we did not detect any significant differences between the two groups with f4 statistics in the form of f4(RMPR_Etruscan, RMPR_Latin; test population, Onge), suggesting shared origins or extensive genetic exchange between them. ... In the Medieval and early modern periods (n = 28 individuals), we observe an ancestry shift toward central and northern Europe in PCA (Fig. 3E), as well as a further increase in the European cluster (C7) and loss of the Near Eastern and eastern Mediterranean clusters (C4 and C5) in ChromoPainter (Fig. 4C). The Medieval population is roughly centered on modern-day central Italians (Fig. 3F). It can be modeled as a two-way combination of Rome's Late Antique population and a European donor population, with potential sources including many ancient and modern populations in central and northern Europe: Lombards from Hungary, Saxons from England, and Vikings from Sweden, among others (table S26). Community commentary: [107] .
  3. ^ Ralph P, Coop G (2013). "The geography of recent genetic ancestry across Europe". PLOS Biology. 11 (5): e1001555. doi:10.1371/journal.pbio.1001555. PMC 3646727. PMID 23667324.
  4. ^ a b c d e Raveane A, Aneli S, Montinaro F, Athanasiadis G, Barlera S, Birolo G, et al. (September 2019). "Population structure of modern-day Italians reveals patterns of ancient and archaic ancestries in Southern Europe". Science Advances. 5 (9): eaaw3492. Bibcode:2019SciA....5.3492R. doi:10.1126/sciadv.aaw3492. PMC 6726452. PMID 31517044.
  5. ^ a b Capocasa M, Anagnostou P, Bachis V, Battaggia C, Bertoncini S, Biondi G, et al. (2014). "Linguistic, geographic and genetic isolation: a collaborative study of Italian populations". Journal of Anthropological Sciences. 92 (92): 201–31. doi:10.4436/JASS.92001. PMID 24607994.
  6. ^ a b c Modi A, Lancioni H, Cardinali I, Capodiferro MR, Rambaldi Migliore N, Hussein A, et al. (July 2020). "The mitogenome portrait of Umbria in Central Italy as depicted by contemporary inhabitants and pre-Roman remains". Scientific Reports. 10 (1): 10700. Bibcode:2020NatSR..1010700M. doi:10.1038/s41598-020-67445-0. PMC 7329865. PMID 32612271.
  7. ^ a b Chiang CW, Marcus JH, Sidore C, Biddanda A, Al-Asadi H, Zoledziewska M, et al. (October 2018). "Genomic history of the Sardinian population". Nature Genetics. 50 (10): 1426–1434. doi:10.1038/s41588-018-0215-8. PMC 6168346. PMID 30224645.
  8. ^ a b c d Marcus JH, Posth C, Ringbauer H, Lai L, Skeates R, Sidore C, et al. (February 2020). "Genetic history from the Middle Neolithic to present on the Mediterranean island of Sardinia". Nature Communications. 11 (1): 939. Bibcode:2020NatCo..11..939M. doi:10.1038/s41467-020-14523-6. PMC 7039977. PMID 32094358.
  9. ^ a b Fernandes DM, Mittnik A, Olalde I, Lazaridis I, Cheronet O, Rohland N, et al. (March 2020). "The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean". Nature Ecology & Evolution. 4 (3): 334–345. Bibcode:2020NatEE...4..334F. doi:10.1038/s41559-020-1102-0. PMC 7080320. PMID 32094539.
  10. ^ a b Sarno, Stefania; Petrilli, Rosalba; Abondio, Paolo; De Giovanni, Andrea; Boattini, Alessio; Sazzini, Marco; De Fanti, Sara; Cilli, Elisabetta; Ciani, Graziella; Gentilini, Davide; Pettener, Davide; Romeo, Giovanni; Giuliani, Cristina; Luiselli, Donata (2021-02-04). "Genetic history of Calabrian Greeks reveals ancient events and long term isolation in the Aspromonte area of Southern Italy". Scientific Reports. 11 (1): 3045. Bibcode:2021NatSR..11.3045S. doi:10.1038/s41598-021-82591-9. ISSN 2045-2322. PMC 7862261. PMID 33542324.
  11. ^ a b Novembre, John; Johnson, Toby; Bryc, Katarzyna; Kutalik, Zoltán; Boyko, Adam R.; Auton, Adam; Indap, Amit; King, Karen S.; Bergmann, Sven; Nelson, Matthew R.; Stephens, Matthew; Bustamante, Carlos D. (2008-08-31). "Genes mirror geography within Europe". Nature. 456 (7218): 98–101. Bibcode:2008Natur.456...98N. doi:10.1038/nature07331. ISSN 0028-0836. PMC 2735096. PMID 18758442.
  12. ^ Tian C, Kosoy R, Nassir R, Lee A, Villoslada P, Klareskog L, et al. (2009). "European population genetic substructure: further definition of ancestry informative markers for distinguishing among diverse European ethnic groups". Molecular Medicine. 15 (11–12): 371–383. doi:10.2119/molmed.2009.00094. PMC 2730349. PMID 19707526.
  13. ^ «Sicily and Southern Italy were heavily colonized by Greeks beginning in the eight to ninth century B.C.. The demographic development of the Greek colonies in Southern Italy was remarkable, and in classical times this region was called Magna Graecia (Great Greece) because it probably surpassed in numbers the Greek population of the motherland.» Cavalli-Sforza L, Menozzi P, Piazza A (1994). The History and Geography of Human Genes. Princeton University Press. p. 278. ISBN 978-0-691-08750-4.
  14. ^ a b c d e Sarno S, Boattini A, Pagani L, Sazzini M, De Fanti S, Quagliariello A, et al. (May 2017). "Ancient and recent admixture layers in Sicily and Southern Italy trace multiple migration routes along the Mediterranean". Scientific Reports. 7 (1): 1984. Bibcode:2017NatSR...7.1984S. doi:10.1038/s41598-017-01802-4. PMC 5434004. PMID 28512355.
  15. ^ Sarno, Stefania; Boattini, Alessio; Pagani, Luca; Sazzini, Marco; De Fanti, Sara; Quagliariello, Andrea; Gnecchi Ruscone, Guido Alberto; Guichard, Etienne; Ciani, Graziella; Bortolini, Eugenio; Barbieri, Chiara; Cilli, Elisabetta; Petrilli, Rosalba; Mikerezi, Ilia; Sineo, Luca (2017-05-16). "Ancient and recent admixture layers in Sicily and Southern Italy trace multiple migration routes along the Mediterranean". Scientific Reports. 7 (1): 1984. Bibcode:2017NatSR...7.1984S. doi:10.1038/s41598-017-01802-4. ISSN 2045-2322. PMC 5434004. PMID 28512355.
  16. ^ Drineas, Petros; Tsetsos, Fotis; Plantinga, Anna; Lazaridis, Iosif; Yannaki, Evangelia; Razou, Anna; Kanaki, Katerina; Michalodimitrakis, Manolis; Perez-Jimenez, Francisco; De Silvestro, Giustina; Renda, Maria C.; Stamatoyannopoulos, John A.; Kidd, Kenneth K.; Browning, Brian L.; Paschou, Peristera (November 2019). "Genetic history of the population of Crete". Annals of Human Genetics. 83 (6): 373–388. doi:10.1111/ahg.12328. ISSN 1469-1809. PMC 6851683. PMID 31192450.
  17. ^ a b c Paschou P, Drineas P, Yannaki E, Razou A, Kanaki K, Tsetsos F, et al. (June 2014). "Maritime route of colonization of Europe". Proceedings of the National Academy of Sciences of the United States of America. 111 (25): 9211–6. Bibcode:2014PNAS..111.9211P. doi:10.1073/pnas.1320811111. PMC 4078858. PMID 24927591.
  18. ^ a b Seldin, Michael F.; Shigeta, Russell; Villoslada, Pablo; Selmi, Carlo; Tuomilehto, Jaakko; Silva, Gabriel; Belmont, John W.; Klareskog, Lars; Gregersen, Peter K. (2006-09-15). "European Population Substructure: Clustering of Northern and Southern Populations". PLOS Genetics. 2 (9): e143. doi:10.1371/journal.pgen.0020143. ISSN 1553-7404. PMC 1564423. PMID 17044734.
  19. ^ a b Stamatoyannopoulos, George; Bose, Aritra; Teodosiadis, Athanasios; Tsetsos, Fotis; Plantinga, Anna; Psatha, Nikoletta; Zogas, Nikos; Yannaki, Evangelia; Zalloua, Pierre; Kidd, Kenneth K.; Browning, Brian L.; Stamatoyannopoulos, John; Paschou, Peristera; Drineas, Petros (8 March 2017). "Genetics of the peloponnesean populations and the theory of extinction of the medieval peloponnesean Greeks". European Journal of Human Genetics. 25 (5): 637–645. doi:10.1038/ejhg.2017.18. ISSN 1476-5438. PMC 5437898. PMID 28272534.
  20. ^ a b Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, et al. (June 2015). "Massive migration from the steppe was a source for Indo-European languages in Europe". Nature. 522 (7555): 207–11. arXiv:1502.02783. Bibcode:2015Natur.522..207H. doi:10.1038/nature14317. PMC 5048219. PMID 25731166.
  21. ^ a b c d Di Gaetano C, Voglino F, Guarrera S, Fiorito G, Rosa F, Di Blasio AM, et al. (2012). "An overview of the genetic structure within the Italian population from genome-wide data". PLOS ONE. 7 (9): e43759. Bibcode:2012PLoSO...743759D. doi:10.1371/journal.pone.0043759. PMC 3440425. PMID 22984441.
  22. ^ a b Price AL, Butler J, Patterson N, Capelli C, Pascali VL, Scarnicci F, et al. (January 2008). "Discerning the ancestry of European Americans in genetic association studies". PLOS Genetics. 4 (1): e236. doi:10.1371/journal.pgen.0030236. PMC 2211542. PMID 18208327.
  23. ^ Nelis M, Esko T, Mägi R, Zimprich F, Zimprich A, Toncheva D, et al. (8 May 2009). "Genetic structure of Europeans: a view from the North-East". PLOS ONE. 4 (5): e5472. Bibcode:2009PLoSO...4.5472N. doi:10.1371/journal.pone.0005472. PMC 2675054. PMID 19424496.
  24. ^ a b c d Fiorito G, Di Gaetano C, Guarrera S, Rosa F, Feldman MW, Piazza A, Matullo G (July 2016). "The Italian genome reflects the history of Europe and the Mediterranean basin". European Journal of Human Genetics. 24 (7): 1056–62. doi:10.1038/ejhg.2015.233. PMC 5070887. PMID 26554880.
  25. ^ a b c Sazzini M, Gnecchi Ruscone GA, Giuliani C, Sarno S, Quagliariello A, De Fanti S, et al. (September 2016). "Complex interplay between neutral and adaptive evolution shaped differential genomic background and disease susceptibility along the Italian peninsula". Scientific Reports. 6: 32513. Bibcode:2016NatSR...632513S. doi:10.1038/srep32513. PMC 5007512. PMID 27582244. Zendo: 165505.
  26. ^ Tian, Chao; Kosoy, Roman; Nassir, Rami; Lee, Annette; Villoslada, Pablo; Klareskog, Lars; Hammarström, Lennart; Garchon, Henri-Jean; Pulver, Ann E.; Ransom, Michael; Gregersen, Peter K.; Seldin, Michael F. (2009). "European population genetic substructure: further definition of ancestry informative markers for distinguishing among diverse European ethnic groups". Molecular Medicine. 15 (11–12): 371–383. doi:10.2119/molmed.2009.00094. ISSN 1528-3658. PMC 2730349. PMID 19707526.
  27. ^ Tian, Chao; Plenge, Robert M.; Ransom, Michael; Lee, Annette; Villoslada, Pablo; Selmi, Carlo; Klareskog, Lars; Pulver, Ann E.; Qi, Lihong; Gregersen, Peter K.; Seldin, Michael F. (January 2008). "Analysis and application of European genetic substructure using 300 K SNP information". PLOS Genetics. 4 (1): e4. doi:10.1371/journal.pgen.0040004. ISSN 1553-7404. PMC 2211544. PMID 18208329.
  28. ^ a b Antonio et al. 2019, p. 2.
  29. ^ a b c Di Gaetano C, Cerutti N, Crobu F, Robino C, Inturri S, Gino S, et al. (January 2009). "Differential Greek and northern African migrations to Sicily are supported by genetic evidence from the Y chromosome". European Journal of Human Genetics. 17 (1): 91–9. doi:10.1038/ejhg.2008.120. PMC 2985948. PMID 18685561. The genetic contribution of Greek chromosomes to the Sicilian gene pool is estimated to be about 37% whereas the contribution of North African populations is estimated to be around 6%.
  30. ^ a b Antonio et al. 2019, p. 3.
  31. ^ Wade, Lizzie (2019). "Immigrants from the Middle East shaped Rome". Science. 366 (6466): 673. Bibcode:2019Sci...366..673W. doi:10.1126/science.366.6466.673. PMID 31699914. S2CID 207965960. Archived from the original on 2021-02-24. Retrieved 2021-01-25."Trade routes sent people and goods to the new capital, and epidemics and invasions reduced Rome's population to about 100,000 people. Invading barbarians brought in more European ancestry. Rome gradually lost its strong genetic link to the Eastern Mediterranean and Middle East. By medieval times, city residents again genetically resembled European populations."
  32. ^ ..."La separazione della Sardegna dal resto del continente, anzi da tutte le altre popolazioni europee, che probabilmente rivela un'origine più antica della sua popolazione, indipendente da quella delle popolazioni italiche e con ascendenze nel Mediterraneo Medio-Orientale." Alberto Piazza, I profili genetici degli italiani, Accademia delle Scienze di Torino
  33. ^ a b c d Brisighelli F, Álvarez-Iglesias V, Fondevila M, Blanco-Verea A, Carracedo A, Pascali VL, et al. (2012). "Uniparental markers of contemporary Italian population reveals details on its pre-Roman heritage". PLOS ONE. 7 (12): e50794. Bibcode:2012PLoSO...750794B. doi:10.1371/journal.pone.0050794. PMC 3519480. PMID 23251386.
  34. ^ Esko T, Mezzavilla M, Nelis M, Borel C, Debniak T, Jakkula E, et al. (June 2013). "Genetic characterization of northeastern Italian population isolates in the context of broader European genetic diversity". European Journal of Human Genetics. 21 (6): 659–65. doi:10.1038/ejhg.2012.229. PMC 3658181. PMID 23249956.
  35. ^ a b Nelis M, Esko T, Mägi R, Zimprich F, Zimprich A, Toncheva D, et al. (2009). "Genetic structure of Europeans: a view from the North-East". PLOS ONE. 4 (5): e5472. Bibcode:2009PLoSO...4.5472N. doi:10.1371/journal.pone.0005472. PMC 2675054. PMID 19424496.
  36. ^ Sazzini M, Abondio P, Sarno S, Gnecchi-Ruscone GA, Ragno M, Giuliani C, et al. (May 2020). "Genomic history of the Italian population recapitulates key evolutionary dynamics of both Continental and Southern Europeans". BMC Biology. 18 (1): 51. doi:10.1186/s12915-020-00778-4. PMC 7243322. PMID 32438927.
  37. ^ Rootsi S (December 2006). "Y-Chromosome haplogroup I prehistoric gene flow in Europe" (PDF). Documenta Praehistorica. 33: 17–20. doi:10.4312/dp.33.3. Archived from the original (PDF) on 2009-03-06.
  38. ^ "Culture del bronzo recente in Italia settentrionale e loro rapporti con la "cultura dei campi di urne"". Gruppo Archeologico Monteclarense. Archived from the original on 10 May 2006.
  39. ^ Haywood J (2014). The Celts: Bronze Age to New Age. Routledge. p. 21. ISBN 978-1-317-87017-3.
  40. ^ "Lombard people". Encyclopaedia Britannica.
  41. ^ Aneli et al. 2022, Supplementary Files:Data S1.
  42. ^ Aneli et al. 2022, pp. 8–11.
  43. ^ A Ciba Foundation Symposium (1959) [1958]. Wolstenholme, Gordon; O'Connor, Cecilia M. (eds.). Medical Biology and Etruscan Origins. London: J & A Churchill Ltd. ISBN 978-0-470-71493-5.
  44. ^ Perkins, Phil (2017). "Chapter 8: DNA and Etruscan identity". In Naso, Alessandro (ed.). Etruscology. Berlin: De Gruyter. pp. 109–118. ISBN 978-1934078495.
  45. ^ Perkins, Phil (2009). "DNA and Etruscan identity". In Perkins, Phil; Swaddling, Judith (eds.). Etruscan by Definition: Papers in Honour of Sybille Haynes. London: The British Museum Research Publications. pp. 95–111. ISBN 978-0861591732. 173.
  46. ^ Reich, David (2018). "Ancient DNA Opens the Floodgates". Who We Are and How We Got Here: Ancient DNA and the New Science of the Human Past. Oxford: Oxford University Press. pp. 53–59. ISBN 9780198821250. But mitochondrial DNA only records information on the entirely female line, a tiny fraction of the many tens of thousands of lineages that have contributed to any person's genome. To understand what really happened in an individual's history, it is incomparably more valuable to examine all ancestral lineages together.
  47. ^ Kron, Geof (2013). "Fleshing out the demography of Etruria". In Macintosh Turfa, Jean (ed.). The Etruscan World. London; New York: Routledge. pp. 56–78. ISBN 978-0-415-67308-2.
  48. ^ a b Perkins, Phil (2017). "DNA and Etruscan identity". In Naso, Alessandro (ed.). Etruscology. Berlin: De Gruyter. pp. 109–18. ISBN 978-1-934078-49-5.
  49. ^ Antonio, Margaret L.; Gao, Ziyue; Moots, Hannah M.; et al. (November 2019). "Ancient Rome: A genetic crossroads of Europe and the Mediterranean". Science. 366 (6466). Washington D.C.: American Association for the Advancement of Science (published November 8, 2019): 708–714. Bibcode:2019Sci...366..708A. doi:10.1126/science.aay6826. hdl:2318/1715466. PMC 7093155. PMID 31699931. Interestingly, although Iron Age individuals were sampled from both Etruscan (n=3) and Latin (n=6) contexts, we did not detect any significant differences between the two groups with f4 statistics in the form of f4(RMPR_Etruscan, RMPR_Latin; test population, Onge), suggesting shared origins or extensive genetic exchange between them.
  50. ^ Posth, Cosimo; Zaro, Valentina; Spyrou, Maria A. (September 24, 2021). "The origin and legacy of the Etruscans through a 2000-year archeogenomic time transect". Science Advances. 7 (39). Washington DC: American Association for the Advancement of Science: eabi7673. Bibcode:2021SciA....7.7673P. doi:10.1126/sciadv.abi7673. PMC 8462907. PMID 34559560.
  51. ^ Bagnasco, G.; Marzullo, M.; Cattaneo, C.; Biehler-Gomez, L.; Mazzarelli, D.; Ricciardi, V.; Müller, W.; Coppa, A.; McLaughlin, R.; Motta, L.; Prato, O.; Schmidt, F.; Gaveriaux, F.; Marras, G. B.; Millet, M. A. (2024-05-28). "Bioarchaeology aids the cultural understanding of six characters in search of their agency (Tarquinia, ninth–seventh century BC, central Italy)". Scientific Reports. 14 (1): 11895. Bibcode:2024NatSR..1411895B. doi:10.1038/s41598-024-61052-z. ISSN 2045-2322. PMC 11133411. PMID 38806487.
  52. ^ Krause, Johannes; Trappe, Thomas (2021) [2019]. A Short History of Humanity: A New History of Old Europe [Die Reise unserer Gene: Eine Geschichte über uns und unsere Vorfahren]. Translated by Waight, Caroline (I ed.). New York: Random House. p. 217. ISBN 9780593229422. It's likely that Basque, Paleo-Sardinian, Minoan, and Etruscan developed on the continent in the course of the Neolithic Revolution. Sadly, the true diversity of the languages that once existed in Europe will never be known.
  53. ^ Antonio et al. 2019, Table 2 Sample Information, Rows 29-32, 36-37.
  54. ^ Antonio et al. 2019, p. 4 Table S15.
  55. ^ Marcus et al. 2020, Supplementary Data 1, A Master Table.
  56. ^ a b Marcus et al. 2020, pp. 5–6.
  57. ^ Marcus et al. 2020, p. 3.
  58. ^ Marcus et al. 2020, pp. 2, 9.
  59. ^ Clemente, Florian; Unterländer, Martina; Dolgova, Olga; Amorim, Carlos Eduardo G.; Coroado-Santos, Francisco; Neuenschwander, Samuel; Ganiatsou, Elissavet; Cruz Dávalos, Diana I.; Anchieri, Lucas; Michaud, Frédéric; Winkelbach, Laura (2021-05-13). "The genomic history of the Aegean palatial civilizations". Cell. 184 (10): 2565–2586.e21. doi:10.1016/j.cell.2021.03.039. ISSN 0092-8674. PMC 8127963. PMID 33930288.
  60. ^ a b Villalba-Mouco, Vanessa; et al. (2021). "Genomic transformation and social organization during the Copper Age–Bronze Age transition in southern Iberia". Science Advances. 7 (47): eabi7038. Bibcode:2021SciA....7.7038V. doi:10.1126/sciadv.abi7038. hdl:10810/54399. PMC 8597998. PMID 34788096.
  61. ^ Manjusha Chintalapati, Nick Patterson, Priya Moorjani (2022) The spatiotemporal patterns of major human admixture events during the European Holocene eLife 11:e77625 https://doi.org/10.7554/eLife.77625
  62. ^ Lai, Luca; Crispu, Stefano (2024), "Social Organization, Intersections, and Interactions in Bronze Age Sardinia. Reading Settlement Patterns in the Area of Sarrala with the Contribution of Applied Sciences", Open Archaeology, 10, doi:10.1515/opar-2022-0358, retrieved 1 September 2024
  63. ^ Serventi, Patrizia; Panicucci, Chiara; Bodega, Roberta; De Fanti, Sara; Sarno, Stefania; Fondevila Alvarez, Manuel; Brisighelli, Francesca; Trombetta, Beniamino; Anagnostou, Paolo; Ferri, Gianmarco; Vazzana, Antonino (2018-01-02). "Iron Age Italic population genetics: the Piceni from Novilara (8th–7th century BC)". Annals of Human Biology. 45 (1): 34–43. doi:10.1080/03014460.2017.1414876. hdl:11573/1085197. ISSN 0301-4460. PMID 29216758.
  64. ^ Cilli, Elisabetta; Cruciani, Fulvio; D’Atanasio, Eugenia; Delpino, Chiara; Finocchi, Stefano; de Gennaro, Luciana; Giacometti, Chiara; Giroldini, Pierluigi; Hajiesmaeil, Mogge; Hui, Ruoyun; Metspalu, Mait; Niinemäe, Helja; Mei, Oscar; Montinaro, Francesco; Pistacchia, Letizia; Ravasini, Francesco; Risi, Flavia; Scheib, Christiana Lyn; Solnik, Anu; Tambets, Kristiina; Trombetta, Beniamino (2024-03-19). "The Genomic portrait of the Picene culture: new insights into the Italic Iron Age and the legacy of the Roman expansion in Central Italy". bioRxiv 10.1101/2024.03.18.585512.
  65. ^ Amorim 2018. "[B]iological relationships played an important role in these early medieval societies... Finally, our data are consistent with the proposed long-distance migration from Pannonia to Northern Italy."
  66. ^ O'Sullivan 2018. "Niederstotzingen North individuals are closely related to northern and eastern European populations, particularly from Lithuania and Iceland."
  67. ^ Vai 2019. "[T]he presence in this cluster of haplogroups that reach high frequency in Northern European populations, suggests a possible link between this core group of individuals and the proposed homeland of different ancient barbarian Germanic groups... This supports the view that the spread of Longobards into Italy actually involved movements of people, who gave a substantial contribution to the gene pool of the resulting populations...This is even more remarkable thinking that, in many studied cases, military invasions are movements of males, and hence do not have consequences at the mtDNA level. Here, instead, we have evidence of maternally linked genetic similarities between LC in Hungary and Italy, supporting the view that immigration from Central Europe involved females as well as males."
  68. ^ Ahlström, Torbjörn; Albris, Sofie Laurine; Allentoft, Morten E.; Altinkaya, Isin; Barrie Andrés, William; Atabiev, Biyaslan; Bangsgaard, Pernille; de Barros Damgaard, Peter; Giovanna Belcastro, Maria; Bergerbrant, Sophie; Bergmann Møller, Stig; Cao, Jialu; Charlier, Nick Card Philippe; Chaume, Bruno; Chernykh, Elizaveta; Trier Christiansen, Torben; Coppa, Alfredo; De Coster, Maura; Demeter, Fabrice; Demoule, Jean-Paul; Dengsø Jessen, Mads; Denham, Sean Dexter; Desenne, Sophie; Downes, Jane; Ellegård, Hanne M.; Ethelberg, Per; Fischer, Anders; Frei, Karin Margarita; Fyfe, Ralph; Gábor, Olivér; Gaillard, Marie-José; Gårdsvoll, Johan Zakarias; Gaunitz, Charleen; Hansen, Jesper; Khaskhanov, Ruslan; Kjær, Katrine; Hedeager, Lotte; Heeren, Stijn; Henning Nielsen, Bjarne; Henriksen, Merete; Henriksen, Rasmus Amund; Heyd, Volker; Høj, Mette; Irving-Pease, Evan K.; Jankauskas, Rimantas; Janson, Henrik; Johansen, Johannsen Torkel; Jørkov, Marie Louise S.; Kähler Holst, Mads; Kern, Anton; Kolosov, Vladimir; Kootker, Lisette M.; Kristiansen, Kristian; Kroonen, Guus; Larsen, Anne Christine; Lejars, Thierry; Løvschal, Mette; Lynnerup, Niels; Magnusson, Yvonne; Mannermaa, Kristiina; Masyakin, Vyacheslav; McColl, Hugh; Melheim, Anne Lene; Merkyte, Inga; Moiseyev, Vyacheslav; Molnár, Erika; Moreno-Mayar, J. Víctor; Morten, Larsen; Mortensen, Fischer; Mortensen, Nadja; Murphy, Eileen; Nielsen, Rasmus; Olsen, Line; Pany-Kucera, Doris; Pinotti, Thomaz; Ponce de León, Marcia S.; Ramsøe, Abigail; Reiersen, Håkon; Reinhard, Walter; Rosengren, Ingason Anders; Sabatini, Serena; Sajantila, Antti; Sand Korneliussen, Thorfinn; Scorrano, Gabriele; Schulz Paulsson, Bettina; Skar, Birgitte; Slavchev, Vladimir; Smrčka, Václav; Staring, Jacqueline; Stenderup, Jesper; Sikora, Martin; Sindbæk, Søren M.; Sørensen, Lasse; Thirup Kastholm, Ole; Tiefengraber, Georg; Tsigaridas Glørstad, Zanette; Uldum, Otto Christian; Valeur Seersholm, Frederik; Vaughn, Andrew; Vega, Jorge; Vimala, Tharsika; Vinner, Lasse; Vitali, Daniele; Voloshinov, Alexey; Wåhlin, Sidsel; Wendling, Holger; Werge, Thomas; Wessman, Anna; Willerslev, Eske; Winther, Jens; Wilhelmson, Helene; Wiltschke, Karin; Yediay, Fulya Eylem; Zilhao, João; Zollikofer, Christoph P. E. (2024-03-04). "Steppe Ancestry in western Eurasia and the spread of the Germanic Languages". bioRxiv 10.1101/2024.03.13.584607.
  69. ^ a b c Grugni V, Raveane A, Mattioli F, Battaglia V, Sala C, Toniolo D, et al. (February 2018). "Reconstructing the genetic history of Italians: new insights from a male (Y-chromosome) perspective". Annals of Human Biology. 45 (1): 44–56. doi:10.1080/03014460.2017.1409801. PMID 29382284. S2CID 43501209.
  70. ^ Di Giacomo F, Luca F, Anagnou N, Ciavarella G, Corbo RM, Cresta M, et al. (September 2003). "Clinal patterns of human Y chromosomal diversity in continental Italy and Greece are dominated by drift and founder effects" (PDF). Molecular Phylogenetics and Evolution. 28 (3): 387–95. Bibcode:2003MolPE..28..387D. doi:10.1016/s1055-7903(03)00016-2. PMID 12927125. Archived from the original (PDF) on 2017-01-20.
  71. ^ Francalacci, P.; Morelli, L.; Underhill, P. A.; Lillie, A. S.; Passarino, G.; Useli, A.; Madeddu, R.; Paoli, G.; Tofanelli, S.; Calò, C. M.; Ghiani, M. E.; Varesi, L.; Memmi, M.; Vona, G.; Lin, A. A. (July 2003). "Peopling of three Mediterranean islands (Corsica, Sardinia, and Sicily) inferred by Y-chromosome biallelic variability". American Journal of Physical Anthropology. 121 (3): 270–279. doi:10.1002/ajpa.10265. ISSN 0002-9483. PMID 12772214.
  72. ^ Francalacci, Paolo; Morelli, Laura; Angius, Andrea; Berutti, Riccardo; Reinier, Frederic; Atzeni, Rossano; Pilu, Rosella; Busonero, Fabio; Maschio, Andrea; Zara, Ilenia; Sanna, Daria; Useli, Antonella; Urru, Maria Francesca; Marcelli, Marco; Cusano, Roberto (2013-08-02). "Low-Pass DNA Sequencing of 1200 Sardinians Reconstructs European Y-Chromosome Phylogeny". Science. 341 (6145): 565–569. Bibcode:2013Sci...341..565F. doi:10.1126/science.1237947. ISSN 0036-8075. PMC 5500864. PMID 23908240.
  73. ^ a b Capelli C, Brisighelli F, Scarnicci F, Arredi B, Caglia' A, Vetrugno G, et al. (July 2007). "Y chromosome genetic variation in the Italian peninsula is clinal and supports an admixture model for the Mesolithic-Neolithic encounter". Molecular Phylogenetics and Evolution. 44 (1): 228–39. Bibcode:2007MolPE..44..228C. doi:10.1016/j.ympev.2006.11.030. PMID 17275346.
  74. ^ Cavalli-Sforza L, Menozzi P, Piazza A (1994). The History and Geography of Human Genes. Princeton University Press. ISBN 978-0-691-08750-4.>: 295 
  75. ^ Semino O, Magri C, Benuzzi G, Lin AA, Al-Zahery N, Battaglia V, et al. (May 2004). "Origin, diffusion, and differentiation of Y-chromosome haplogroups E and J: inferences on the neolithization of Europe and later migratory events in the Mediterranean area". American Journal of Human Genetics. 74 (5): 1023–34. doi:10.1086/386295. PMC 1181965. PMID 15069642.
  76. ^ Barker, Graeme; Rasmussen, Tom (2000). The Etruscans. The Peoples of Europe. Oxford: Blackwell Publishing. p. 44. ISBN 978-0-631-22038-1.
  77. ^ Turfa, Jean MacIntosh (2017). "The Etruscans". In Farney, Gary D.; Bradley, Gary (eds.). The Peoples of Ancient Italy. Berlin: De Gruyter. pp. 637–672. doi:10.1515/9781614513001. ISBN 978-1-61451-520-3.
  78. ^ De Grummond, Nancy T. (2014). "Ethnicity and the Etruscans". In McInerney, Jeremy (ed.). A Companion to Ethnicity in the Ancient Mediterranean. Chichester, UK: John Wiley & Sons, Inc. pp. 405–422. doi:10.1002/9781118834312. ISBN 9781444337341.
  79. ^ Shipley, Lucy (2017). "Where is home?". The Etruscans: Lost Civilizations. London: Reaktion Books. pp. 28–46. ISBN 9781780238623.
  80. ^ a b Messina F, Finocchio A, Rolfo MF, De Angelis F, Rapone C, Coletta M, et al. (2015). "Traces of forgotten historical events in mountain communities in Central Italy: A genetic insight". American Journal of Human Biology. 27 (4): 508–19. doi:10.1002/ajhb.22677. PMID 25728801. S2CID 30111156.
  81. ^ Capelli C, Onofri V, Brisighelli F, Boschi I, Scarnicci F, Masullo M, et al. (June 2009). "Moors and Saracens in Europe: estimating the medieval North African male legacy in southern Europe". European Journal of Human Genetics. 17 (6): 848–52. doi:10.1038/ejhg.2008.258. PMC 2947089. PMID 19156170.
  82. ^ a b c d Sarno S, Boattini A, Carta M, Ferri G, Alù M, Yao DY, et al. (2014). "An ancient Mediterranean melting pot: investigating the uniparental genetic structure and population history of sicily and southern Italy". PLOS ONE. 9 (4): e96074. Bibcode:2014PLoSO...996074S. doi:10.1371/journal.pone.0096074. PMC 4005757. PMID 24788788. This article contains quotations from this source, which is available under a Creative Commons Attribution 4.0 International (CC BY 4.0) license.
  83. ^ Brisighelli, F (2012). "Uniparental markers of contemporary Italian population reveals details on its pre-Roman heritage". PLOS ONE. 7 (12): e50794. Bibcode:2012PLoSO...750794B. doi:10.1371/journal.pone.0050794. PMC 3519480. PMID 23251386. In Table S4, #BEL50 is estimated to be Q-M378 by haplotype, though it is shown as just P* (xR1).
  84. ^ a b Pereira L, Richards M, Goios A, Alonso A, Albarrán C, Garcia O, et al. (January 2005). "High-resolution mtDNA evidence for the late-glacial resettlement of Europe from an Iberian refugium". Genome Research. 15 (1): 19–24. doi:10.1101/gr.3182305. PMC 540273. PMID 15632086.
  85. ^ Richards M, Macaulay V, Hickey E, Vega E, Sykes B, Guida V, et al. (November 2000). "Tracing European founder lineages in the Near Eastern mtDNA pool". American Journal of Human Genetics. 67 (5): 1251–76. doi:10.1016/S0002-9297(07)62954-1. PMC 1288566. PMID 11032788.
  86. ^ Achilli A, Rengo C, Magri C, Battaglia V, Olivieri A, Scozzari R, et al. (November 2004). "The molecular dissection of mtDNA haplogroup H confirms that the Franco-Cantabrian glacial refuge was a major source for the European gene pool". American Journal of Human Genetics. 75 (5): 910–8. doi:10.1086/425590. PMC 1182122. PMID 15382008.
  87. ^ Achilli A, Olivieri A, Pala M, Metspalu E, Fornarino S, Battaglia V, et al. (April 2007). "Mitochondrial DNA variation of modern Tuscans supports the near eastern origin of Etruscans". American Journal of Human Genetics. 80 (4): 759–68. doi:10.1086/512822. PMC 1852723. PMID 17357081. 4/138=2.9% in Latium; 3/114=2.6% in Volterra; 2/92=2.2% in Basilicata and 3/154=2.0% in Sicily
  88. ^ Boattini A, Martinez-Cruz B, Sarno S, Harmant C, Useli A, Sanz P, et al. (2013). "Uniparental markers in Italy reveal a sex-biased genetic structure and different historical strata". PLOS ONE. 8 (5): e65441. Bibcode:2013PLoSO...865441B. doi:10.1371/journal.pone.0065441. PMC 3666984. PMID 23734255.
  89. ^ Balter M (3 June 2010). "Tracing the Roots of Jewishness". Science Magazine. American Association for the Advancement of Science.
  90. ^ Zoossmann-Diskin A (October 2010). "The origin of Eastern European Jews revealed by autosomal, sex chromosomal and mtDNA polymorphisms". Biology Direct. 5 (57): 57. doi:10.1186/1745-6150-5-57. PMC 2964539. PMID 20925954.
  91. ^ Balter M (8 October 2013). "Did Modern Jews Originate in Italy?". ScienceNOW.
  92. ^ Yandell K (2013). "Genetic Roots of the Ashkenazi Jews". The Scientist.
  93. ^ Rosenberg NA, Pritchard JK, Weber JL, Cann HM, Kidd KK, Zhivotovsky LA, Feldman MW (December 2002). "Genetic structure of human populations". Science. 298 (5602): 2381–5. Bibcode:2002Sci...298.2381R. doi:10.1126/science.1078311. PMID 12493913. S2CID 8127224.
  94. ^ Costa MD, Pereira JB, Pala M, Fernandes V, Olivieri A, Achilli A, et al. (2013). "A substantial prehistoric European ancestry amongst Ashkenazi maternal lineages". Nature Communications. 4: 2543. Bibcode:2013NatCo...4.2543C. doi:10.1038/ncomms3543. PMC 3806353. PMID 24104924.
  95. ^ Banda Y, Kvale M, Hoffmann T, Hesselson S, Tang H, Ranatunga D, et al. (2013). "Admixture Estimation in a Founder Population". Am Soc Hum Genet. Archived from the original on 2019-08-11. Retrieved 2016-10-31.
  96. ^ Bray SM, Mulle JG, Dodd AF, Pulver AE, Wooding S, Warren ST (September 2010). "Signatures of founder effects, admixture, and selection in the Ashkenazi Jewish population". Proceedings of the National Academy of Sciences of the United States of America. 107 (37): 16222–7. Bibcode:2010PNAS..10716222B. doi:10.1073/pnas.1004381107. PMC 2941333. PMID 20798349.
  97. ^ Atzmon G, Hao L, Pe'er I, Velez C, Pearlman A, Palamara PF, Morrow B, Friedman E, Oddoux C, Burns E, Ostrer H (June 2010). "Abraham's children in the genome era: major Jewish diaspora populations comprise distinct genetic clusters with shared Middle Eastern Ancestry". American Journal of Human Genetics. 86 (6): 850–9. doi:10.1016/j.ajhg.2010.04.015. PMC 3032072. PMID 20560205.
  98. ^ "Genes Set Jews Apart, Study Finds". American Scientist. Retrieved 8 November 2013.
  99. ^ Pala M, Achilli A, Olivieri A, Hooshiar Kashani B, Perego UA, Sanna D, et al. (June 2009). "Mitochondrial haplogroup U5b3: a distant echo of the epipaleolithic in Italy and the legacy of the early Sardinians" (PDF). American Journal of Human Genetics. 84 (6): 814–21. doi:10.1016/j.ajhg.2009.05.004. PMC 2694970. PMID 19500771. Archived from the original (PDF) on 2018-07-22. Retrieved 2010-07-15.
  100. ^ Leonardi, Michela; Sandionigi, Anna; Conzato, Annalisa; Vai, Stefania; Lari, Martina (2018). "The female ancestor's tale: Long-term matrilineal continuity in a nonisolated region of Tuscany". American Journal of Physical Anthropology. 167 (3). New York City: John Wiley & Sons (published September 6, 2018): 497–506. doi:10.1002/ajpa.23679. PMID 30187463. S2CID 52161000.
  101. ^ Kovacevic L, Tambets K, Ilumäe AM, Kushniarevich A, Yunusbayev B, Solnik A, et al. (2014). "Standing at the gateway to Europe--the genetic structure of Western balkan populations based on autosomal and haploid markers". PLOS ONE. 9 (8): e105090. Bibcode:2014PLoSO...9j5090K. doi:10.1371/journal.pone.0105090. PMC 4141785. PMID 25148043.
  102. ^ Wade N (13 August 2008). "Genetic Map of Europe". New York Times. Retrieved 25 February 2014.
  103. ^ Li JZ, Absher DM, Tang H, Southwick AM, Casto AM, Ramachandran S, Cann HM, Barsh GS, Feldman M, Cavalli-Sforza LL, Myers RM (February 2008). "Worldwide human relationships inferred from genome-wide patterns of variation" (PDF). Science. 319 (5866). New York, N.Y.: 1100–4. Bibcode:2008Sci...319.1100L. doi:10.1126/science.1153717. PMID 18292342. S2CID 53541133. Archived from the original (PDF) on 2009-03-27.
  104. ^ López Herráez D, Bauchet M, Tang K, Theunert C, Pugach I, Li J, et al. (November 2009). "Genetic variation and recent positive selection in worldwide human populations: evidence from nearly 1 million SNPs". PLOS ONE. 4 (11): e7888. Bibcode:2009PLoSO...4.7888L. doi:10.1371/journal.pone.0007888. PMC 2775638. PMID 19924308.
  105. ^ Moorjani P, Patterson N, Hirschhorn JN, Keinan A, Hao L, Atzmon G, Burns E, Ostrer H, Price AL, Reich D (April 2011). "The history of African gene flow into Southern Europeans, Levantines, and Jews". PLOS Genetics. 7 (4): e1001373. doi:10.1371/journal.pgen.1001373. PMC 3080861. PMID 21533020.
  106. ^ Keller A, Graefen A, Ball M, Matzas M, Boisguerin V, Maixner F, et al. (February 2012). "New insights into the Tyrolean Iceman's origin and phenotype as inferred by whole-genome sequencing". Nature Communications. 3: 698. Bibcode:2012NatCo...3..698K. doi:10.1038/ncomms1701. PMID 22426219.
  107. ^ Provyn, Hunter (2019-11-08). "First Ancient J-Z631 Found in the Via Salaria Necropolis in Rome". Phylogeographer.
  108. ^ Raveane, Alessandro; Molinaro, Ludovica; Aneli, Serena; Capodiferro, Marco Rosario; Ongaro, Linda; Migliore, Nicola Rambaldi; Soffiati, Sara; Scarano, Teodoro; Torroni, Antonio; Achilli, Alessandro; Ventura, Mario; Pagani, Luca; Capelli, Cristian; Olivieri, Anna; Bertolini, Francesco (2022-03-01). "Assessing temporal and geographic contacts across the Adriatic Sea through the analysis of genome-wide data from Southern Italy". bioRxiv: 2022.02.26.482072. doi:10.1101/2022.02.26.482072. hdl:11586/436540. S2CID 247231413.
  109. ^ Balvanović, R.; Milović, O.; Spasić-Đurić, D.; Stojanović, M. M.; Šmit, Ž.; Špehar, P. (2022-04-02) [received 2021-10-27]. "Late Roman glass from Viminacium and Egeta (Serbia): glass-trading patterns on Iron Gates Danubian Limes". Archaeological and Anthropological Sciences. 14 (79). doi:10.1007/s12520-022-01529-y. ISSN 1866-9557.
  110. ^ Various (2021-08-31). "Тема: дДНК из Сербии от римского времени до средних веков". Молекулярная генеалогия.
  111. ^ Various (2021-08-31). "Genetic study: Ancient DNA of Roman Danubian Frontier and Slavic Migrations (Olalde 2021)". Eupedia.com.
  112. ^ Ambron (2021-08-31). "Cosmopolitanism at the Roman Danubian Frontier, Slavic Migrations, and the Genomic Formation of Modern Balkan Peoples". Slavic origins.
  113. ^ Khan, Razib (2021-08-31). "The genetic history of the Serbs (and proto-Serbs)". Gene Expression.
  114. ^ Various (2021-09-01). "Cosmopolitanism at the Roman Danubian Frontier, Slavic Migrations, and the Genomic Formation of Modern Balkan Peoples". Anthrogenica.
  115. ^ Various (2021-09-01). "Одг: Древна ДНК - научни радови". Порекло.
  116. ^ Sécher, Bernard (2021-09-03). "Le cosmopolitisme à la frontière Romaine danubienne". Généalogie génétique.
  117. ^ Weselowski, David (2021-09-04). "The genomic formation of modern Balkan peoples (Olalde et al. 2021 preprint)". Eurogenes Blog: Focusing on ancient population genomics.
  118. ^ Andvari (2021-09-09). "Как римляне и славяне влияли на генетику Балкан". Станция утешения: Люблю первобытное общество и все, что с ним связано.
  119. ^ Oblakov-Drim, Ivan (2023-04-01). "Балто-славяне в римском Виминациуме". Проза.ру.
  120. ^ Васильченко, Валерий (2023-04-08). "Генетика о славянизации Балкан". Дзен.ру.
  121. ^ Various (2023-12-07). "A genetic history of the Balkans from Roman frontier to Slavic migrations". GenArchivist: Populations and Genetics Forums.
  122. ^ Various (2023-12-07). "Тема: A genetic history of the Balkans from Roman frontier to Slavic migrations". Порекло.
  123. ^ Various (2023-12-07). "Genetic study: A genetic history of the Balkans from Roman frontier to Slavic migrations". Eupedia.com.
  124. ^ A Cypriot (2023-12-10). "Did ancient Greeks leave a genetic impact on West Anatolia? qpAdm and G25 analysis". Genes of the Ancients. Archived from the original on 2024-04-27.
  125. ^ Bading, Ingo (2023-12-15). "An der mittleren Donau - Die Goten sterben aus, unter den Awaren kommt es zur Zuwanderung der Slawen: Sind Goten und Sarmaten auf dem Balkan genetisch ausgestorben?". Studium generale: Wissenschaftsblog - Kosmos, Erde, Geschichte. Archived from the original on 2023-12-22.
  126. ^ "A genetic history of the Balkans from Roman frontier to Slavic migrations". PlumX Metrics. 2023-12-07.
  127. ^ Olalde; et al. (2023-12-07) [received 2023-06-23]. "A genetic history of the Balkans from Roman frontier to Slavic migrations". Cell. 186 (25): 5472–5485. eISSN 1097-4172. Preprint published as Olalde; et al. (2021-09-04). "Cosmopolitanism at the Roman Danubian Frontier, Slavic Migrations, and the Genomic Formation of Modern Balkan Peoples". bioRxiv 10.1101/2021.08.30.458211v1. Academic commentary: [109] . Community commentary: [110]˙​[111]˙​[112][113]˙​[114]˙​[115]˙​[116]˙​[117]˙​[118]˙​[119]˙​[120]˙​[121]˙​[122]˙​[123]˙​[124]˙​.[125] News: [126] .
  128. ^ Monnereau, Aurore; Ughi, Alice; Orecchioni, Paola; Hagan, Richard; Talbot, Helen M.; Nikita, Efthymia; Hamilton, Derek; Le Roux, Petrus; Molinari, Alessandra; Carver, Martin; Craig, Oliver E.; Speller, Camilla F.; Alexander, Michelle M.; Wales, Nathan (24 July 2024). "Multi-proxy bioarchaeological analysis of skeletal remains shows genetic discontinuity in a Medieval Sicilian community". Royal Society Open Science. 11 (7). Bibcode:2024RSOS...1140436M. doi:10.1098/rsos.240436. ISSN 2054-5703. PMC 11265863. PMID 39050717.
  129. ^ Yeshurun, Reuven (March 2024). "Signalling Palaeolithic identity". Nature Human Behaviour. 8 (published 2024-01-29): 414–415. doi:10.1038/s41562-023-01805-4. eISSN 2397-3374.
  130. ^ Гаджиева, Алиса (2024-01-30). "Доисторическая бижутерия разделила граветтскую культуру". Naked Science.
  131. ^ Wild, Sarah (2024-01-29). "Ancient Jewelry Shows Ice Age Europe Had 9 Distinct Cultures". Scientific American. ISSN 0036-8733.
  132. ^ Nalewicki, Jennifer (2024-01-29). "Prehistoric jewelry reveals 9 distinct cultures across Stone Age Europe". Live Science.
  133. ^ Guesgen, Mirjam (2024-01-30). "Scientists Discover Stunning Evidence of Multiple Lost Prehistoric Societies: Researchers analyzed more than 100 pieces of prehistoric jewelry and found that the ancient past was more complex than we imagined". Vice. ISSN 1077-6788.
  134. ^ Yirka, Bob (2024-01-30). "Study of ancient adornments suggests nine distinct cultures lived in Europe during the Paleolithic". Phys.org.
  135. ^ Watson, Clare (2024-01-31). "Stunning Prehistoric Jewelry Reveals 9 Hidden Societies in Ice Age Europe". ScienceAlert.
  136. ^ Kamrani, Kambiz (2024-02-01). "Unlocking Ancient Secrets: Jewelry Reveals Nine Lost Cultures: Decoding Human Histories Through Prehistoric Adornments". Anthropology.net.
  137. ^ Baker, Jack; Courtenay, Lloyd A.; d’Errico, Francesco; Rigaud, Solange; Pereira, Daniel (2024-01-29) [received 2023-04-26]. "Evidence from personal ornaments suggest nine distinct cultural groups between 34,000 and 24,000 years ago in Europe". Nature Human Behaviour. 8: 431–444. doi:10.1038/s41562-023-01803-6. eISSN 2397-3374. Academic commentary: [129] . Community commentary: [130] . News: [131]˙​[132]˙​[133]˙​[134]˙​[135]˙​.[136]
  138. ^ Globe Institute (2023-09-21). "Project PRJEB64656: Population Genomics of Late Stone Age Western Eurasia". European Nucleotide Archive.
  139. ^ Allentoft, Morten E.; Sikora, Martin; Willerslev, Eske (2023-09-22). "Genotype data from Population Genomics of Postglacial Western Eurasia". University of Copenhagen Electronic Research Data Archive.
  140. ^ Allentoft, Morten E.; Sikora, Martin; Willerslev, Eske (2023-09-22). "Supporting data for "Population Genomics of Postglacial Western Eurasia"". Zenodo.
  141. ^ Attwaters, Michael (March 2024). "Ancient migration and the modern genome". Nature Reviews Genetics. 25 (162) (published 2024-02-05). doi:10.1038/s41576-024-00702-4. eISSN 1471-0064.
  142. ^ Alexandersen, Verner; Allentoft, Morten E.; Bennike, Ole; Gotfredsen, Anne Birgitte; Gröcke, Darren R.; Fischer, Anders; Ingason, Andrés; Iversen, Rune; Jørkov, Marie Louise; Kristiansen, Kristian; Lysdahl, Per; Refoyo-Martínez, Alba; Price, T. Douglas; Racimo, Fernando; Scorrano, Gabriele; Sikora, Martin; Sjögren, Karl-Göran; Sørensen, Lasse; Stenderup, Jesper; Zetner Trolle Jensen, Theis; Wåhlin, Sidsel; Vimala, Tharsika; Willerslev, Eske (2024-02-14). "Vittrup Man–The life-history of a genetic foreigner in Neolithic Denmark". Public Library of Science One. eISSN 1938-1352.
  143. ^ Khan, Razib (2022-05-05). "Eurasia, the Stone Age and revenge of the Danes!". Gene Expression.
  144. ^ Various (2022-05-06). "Massive new paper from Allenthoft and Willerslev". Anthrogenica.
  145. ^ Oh-Willeke, Andrew (2022-05-06). "A Eurasian Stone Age Ancient DNA Megapaper". Dispatches From Turtle Island: Observations That Transcend Law and Politics.
  146. ^ Various (2023-05-06). "RE: The story of the Khvalynsk people". Eurogenes Blog: Focusing on ancient population genomics.
  147. ^ Wesolowski, David (2022-05-07). "Population genomics of Stone Age Eurasia (Allentoft et al. 2022 preprint)". Eurogenes Blog: Focusing on ancient population genomics.
  148. ^ Stone Age Herbalist (2022-05-09). "Breakdown - 'Population Genomics Of Stone Age Eurasia' Paper: New Genetics Results Covering The Mesolithic-Neolithic Transition". Grey Goose Chronicles.
  149. ^ Sécher, Bernard (2022-05-10). "L'histoire génomique de l'Eurasie à l'Âge de Pierre". Généalogie génétique.
  150. ^ Andvari (2022-05-15). "Популяционная генетика каменного века Евразии". Станция утешения: Люблю первобытное общество и все, что с ним связано.
  151. ^ Various (2023-09-29). "Тема: Allentoft et al: Population Genomics of Late Stone Age West Eurasia". Порекло.
  152. ^ Various (2023-09-29). "Allentoft Population Genomics of Late Stone Age West Eurasia". GenArchivist: Populations and Genetics Forum.
  153. ^ Various (2023-10-22). "RE: The Origin of R1b-P312". GenArchivist.
  154. ^ Подрезов, Михаил (2024-01-10). "Палеогенетики нашли предков ямников на Среднем Дону: Этот один из результатов исследования 317 древних геномов". N + 1 Интернет-издание.
  155. ^ Various (2024-01-13). "RE: The Origin of R1b-P312 Bell Beaker". GenArchivist: Populations and Genetics Forum.
  156. ^ Various (2024-01-12). "Time Maps main post-glacial European ancestry components". GenArchivist: Populations and Genetics Forum.
  157. ^ "RE: aDNA (Dodecad K12b)". Eupedia. 2024-03-15.
  158. ^ Khan, Razib (2024-03-28). "Kristian Kristiansen: DNA and European prehistory: One of the preeminent archaeologists of Northern Europe talks about where we are in 2024". Razib Khan's Unsupervised Learning.
  159. ^ Hillgren, Johanna (2024-01-10). "New light shed on the Stone Age 'invisible wall'". University of Gothenburg.
  160. ^ Department of Environmental Biology Nature (2024-01-12). "The invisible genetic barrier that divided western Eurasia in prehistory". Sapienza Università di Roma.
  161. ^ Schuster, Ruth (2024-01-16). "Who Are Western Europeans? New Study Reveals True Origins". Haaretz.
  162. ^ Kamrani, Kambiz (2024-01-20). "Unraveling European Genomic Mysteries: Ancient DNA and the Legacy of Migration". Anthropology.net.
  163. ^ Gaglioti, Frank (2024-03-21). "Genetic makeup of modern Europeans derived through ancient migrations from Asia". World Socialist Web Site.
  164. ^ "Population genomics of post-glacial western Eurasia". Altmetric. 2024-01-10.
  165. ^ Allentoft, Morten E.; Altinkaya, Isin; et al. (2024-01-11) [received 2022-05-02]. "Population genomics of post-glacial western Eurasia". Nature (625) (published 2024-01-10): 301–311. eISSN 1476-4687. Preprint first published as "Population Genomics of Stone Age Eurasia". 2022-05-05. bioRxiv 10.1101/2022.05.04.490594v1. Data: [138]˙​[139]˙​.[140] Academic commentary: [141]˙​.[142] Community commentary: [143]˙​[144]˙​[145]˙​[146]˙​[147]˙​[148]˙​[149]˙​[150]˙​[151]˙​[152]˙​[153]˙​[154]˙​[155]˙​[156]˙​[157]˙​.[158] News: [159]˙​[160]˙​[161]˙​[162]˙​[163]˙​.[164]
  166. ^ Uppsala University (2024-01-15). "Project PRJEB71770: Analyses of complete genomes of 10 French Late Mesolithic individuals form the sites of Hoedic, Téviec and Champigny". European Nucleotide Archive.
  167. ^ Rowley-Conwy, Peter (2024-02-26). "Hunter-gatherers and earliest farmers in western Europe". Proceedings of the National Academy of Sciences. 121 (10). doi:10.1073/pnas.2322683121. eISSN 1091-6490. PMC 10927556.
  168. ^ Various (2024-01-18). "Analyses of complete genomes of 10 French Late Mesolithic individuals form the sites". GenArchivist: Populations and Genetics Forum.
  169. ^ Various (2024-01-24). "Тема: Порекло хаплогрупе I2-Y3120 у светлу нових археогенетичких сазнања". Порекло.
  170. ^ Парфенов, Василий (2024-03-02). "Анализ генома охотников-собирателей показал, как в каменном веке избегали кровосмешения". Naked Science.
  171. ^ Sécher, Bernard (2024-03-02). "Génomes anciens des derniers chasseurs-cueilleurs de France". Généalogie génétique.
  172. ^ Uppsala University Press and Media Services (2024-02-28). "Stone Age strategy for avoiding inbreeding". Uppsala Universitet.
  173. ^ Carvajal, Guillermo (2024-02-28). "Researchers Reveal how Stone Age Hunter-Gatherers Avoided Inbreeding". La Brújula Verde: Magazine Cultural Independiente.
  174. ^ Kamrani, Kambiz (2024-03-07). "Unraveling Social Dynamics: Ancient DNA Sheds Light on Europe's Last Hunter-Gatherers: Insights into Cultural Strategies and Social Bonds Amidst Neolithic Transition". Anthropology.net.
  175. ^ Gutierrez C., Nicolas (2024-03-22). "Comment les derniers chasseurs-cueilleurs de France évitaient la consanguinité". Sciences et Avenir. ISSN 0036-8636.
  176. ^ Capone, Franco (2024-03-27). "I cacciatori raccoglitori della Bretagna non praticavano l'incesto, anche a rischio di "estinguersi": Gli ultimi cacciatori raccoglitori europei non si incrociavano con gli agricoltori ed evitavano i rapporti incestuosi, anche se erano rimasti in pochi". Focus.
  177. ^ Alaçamlı, Erkin; Bernhardsson, Carolina; Bouquin, Denis; Chetty, Darshan; Dina, Christian; Edlund, Hanna; Garmond, Nicolas; Günther, Torsten; Jakobsson, Mattias; Marchand, Grégor; Simões, Luciana G.; Peyroteo-Stjerna, Rita (2024-02-26). "Genomic ancestry and social dynamics of the last hunter-gatherers of Atlantic France". Proceedings of the National Academy of Sciences. 121 (10). doi:10.1073/pnas.2310545121. eISSN 1091-6490. Data: [166] . Academic commentary: [167] . Community commentary: [168]˙​[169]˙​[170]˙​.[171] News: [172]˙​[173]˙​[174]˙​[175]˙​.[176]
  178. ^ Ankerl, Barbara; Cheronet, Olivia; Greisinger, Michaela; Hagmann, Dominik; Kirchengast, Nisa Iduna; Kirchengast, Sylvia; Miglbauer, Renate (March 2024) [2023-12-02]. "Double feature: First genetic evidence of a mother-daughter double burial in Roman period Austria". Journal of Archaeological Science: Reports. 55 (published May 2024). ISSN 2352-409X.
  179. ^ Various (2024-02-15). "RE: Genetic Genealogy & Ancient DNA (DISCUSSION ONLY)". GenArchivist.
  180. ^ Various (2024-02-18). "Until death do us part". Eupedia.
  181. ^ Various (2024-02-19). "The Late Iron Age Celtic Necropolis of Seminario Vescovile in Verona, Italy". FamilyTreeDNA.
  182. ^ Various (2024-02-20). "Iron Age R1b-L21 from the Celtic Cenomani Tribe in Northern Italy". GenArchivist.
  183. ^ Various (2024-03-06). "RE: Ancient I1 samples list and discussion". GenArchivist.
  184. ^ Various (2024-03-07). "An I-M253 among the Cisalpine Celts?". GenArchivist.
  185. ^ Walters, Sam (2024-02-14). "Ancient Italians' Affection for Animals Went Deep, All the Way to Their Graves". Discover Magazine. ISSN 0274-7529.
  186. ^ Carvajal, Guillermo (2024-02-18). "Unknown Celtic Rituals and Beliefs May Explain Strange Burials of People with Animal Parts Found in Verona". La Brújula Verde: Magazine Cultural Independiente.
  187. ^ Amato, Alfonsina; Arenz, Gabriele; Bersani, Marzia; Coia, Valentina; Dori, Irene; Laffranchi, Zita; Lösch, Sandra; Milella, Marco; Paladin, Alice; Ryan-Despraz, Jessica; Salzani, Luciano; Szidat, Sönke; Tecchiati, Umberto; Thompson, Simon R.; Zingale, Stefania; Zink, Albert (2024-02-14). ""Until death do us part": A multidisciplinary study on human- Animal co- burials from the Late Iron Age necropolis of Seminario Vescovile in Verona (Northern Italy, 3rd-1st c. BCE)". Public Library of Science One. eISSN 1938-1352. Academic commentary: [178] . Community commentary: [179]˙​[180]˙​[181]˙​[182]˙​[183]˙​.[184] News: [185]˙​.[186]

Sources

[edit]

Further reading

[edit]
  • Saupe, Tina et al. "Ancient genomes reveal structural shifts after the arrival of Steppe-related ancestry in the Italian Peninsula". In: Current Biology Volume 31, Issue 12, 21 June 2021, Pages 2576–2591.e12. DOI: https://doi.org/10.1016/j.cub.2021.04.022
Retrieved from "https://en.wikipedia.org/w/index.php?title=Genetic_history_of_Italy&oldid=1276768433"