African admixture in Europe

Source: Wikipedia, the free encyclopedia.
For the African ancestry of all non-African Paleolithic populations, see Recent African origin of modern humans and Southern Dispersal.

African admixture in Europe refers to the presence of human genotypes attributable to periods of human population dispersals out of Africa in the genetic history of Europe. For example, certain Y-DNA and mtDNA lineages are thought to have spread from Northeastern Africa to the Near East during the later Pleistocene, and from there to Europe with the Neolithic Revolution.[1][2]

More recent African admixture – primarily

Iberian peninsula (modern day Spain and Portugal), with higher levels in the West and the South[3][4][5] and Southern Italy, with higher levels in Sardinia and Sicily.[6]

Neolithic

The change from

Asia Minor around 10,000 BCE. Agriculture was introduced into Europe and North Africa by migrating farmers from West Asia.[7] According to the demic diffusion model, these Middle Eastern farmers either replaced or interbred with the local hunter-gather populations that had been living in Europe since the Out of Africa migration.[8]

It has been suggested that the first Middle Eastern farmers reflected North African influences or vice versa.[9] There have been suggestions that some genetic lineages found in the Middle East arrived there during this period.[10] The first agricultural societies in the Middle East are generally thought to have emerged after, and perhaps from, the Natufian culture between 12,000 and 10,000 BCE. The latter group was widely semi-sedentary even before the introduction of agriculture. An important migration from North Africa across the Sinai also appears to have occurred before the formation of the Natufian.[citation needed].

Historical period

In historical times, there has been a period of north African influence in southern Europe, especially in the

colonization of Africa and as a consequence of the Atlantic slave trade.[2]

Admixture

5 component admixture plots for European, West Asian, North African and West African populations (Hernández et al. 2019)[11]
  • Hernandez et al. (2020) identified 11.17 ± 1.87% North African ancestry in southern Portuguese samples (from a population similar to modern northern Moroccans and Algerians), 9.28 ± 1.79% of such ancestry in western Andalusians, and an average of 1.41 ± 0.72% sub-Saharan ancestry in southern Iberians (using Yoruba as a proxy source). Substantially lower levels of North African admixture were further detected in Northern Italians (0.77%) and Tuscans (1%). [11]
  • Olalde et al. (2019) found evidence for 'sporadic contacts' between Iberia and North Africa in the
    Copper Age and Bronze Age. A male sample from central Iberia, dating from 2473–2030 cal BCE, was found to cluster with modern and ancient North Africans, characterised by ancestry from both Late Pleistocene North Africans and Early Neolithic Europeans. Another Bronze Age sample had 25% such North African ancestry. However, North African ancestry only became widespread in Iberia in the past ~2000 years, associated with the Roman Empire or earlier Punic presence and the later period of Muslim rule. The study analysed 45 samples from southeastern Spain dating from the 3rd-16th centuries CE, all of which fell outside the genetic variation of preceding Iberian Iron Age populations, harbouring ancestry from both southern European and North African populations, as well as additional Levantine-related ancestry. 2 samples out of 23 dating from the 10th to 16th centuries were also found to have partial sub-Saharan ancestry, which was not identified in earlier samples. Present-day southern Iberians harbour less African ancestry than Muslim period samples, likely due to subsequent population expulsions and repopulation from the north, as supported by historical sources and genetic analysis of present-day groups.[4]
Distribution of North African Admixture in the Iberian Peninsula (Bycroft et al. 2019)[5]
  • Bycroft et al. (2019) identified regionally varying fractions of Northwest African ancestry in modern Iberians, ranging from 0–12%. This ancestry was found to be from a source population similar to modern Northwest Africans. The admixture was dated to 860–1120 CE, associated with the Muslim conquest and subsequent Reconquista. The highest levels of Northwest African admixture were identified in western Iberia whilst the lowest levels were found in the Basque region and an area in the North East roughly corresponding to the 14th-century Crown of Aragon. They also found some evidence for a second admixture event in Portuguese and Southern Spanish groups involving a second North African population within which a small sub-Saharan African component was detected.This admixture event was dated to approximately 1300 CE.[5]
Haplotype-based estimates of genetic sharing between Europe and Africa (Botigué et al. 2013)[12]
  • Botigué et al. (2013) analysed genome-wide SNP data from over 2,000 modern individuals from Iberia. They estimated an average of 4% to 12% Northwest African admixture in modern Iberians (with low or zero levels in Basques), whereas populations in southeastern Europe had less than 2% of such admixture. Sub-Saharan African ancestry was detected at less than 1% in Europe, with the exception of the Canary Islands.[12][13]
  • Moorjani et al. (2011) estimated that some Southern Europeans have inherited 1%–3% sub-Saharan ancestry (approximately 3.2% in Portugal, 2.9% in Sardinia, 2.7% in southern Italy, 2.4% in Spain and 1.1% in northern Italy), although the percentages were lower (ranging from 0.2% in Sardinia and northern Italy to 2.1% in Portugal) when reanalyzed with the 'STRUCTURE' statistical model. An average mixture date of around 55 generations/1100 years ago was given, "consistent with North African gene flow at the end of the Roman Empire and subsequent Arab migrations". This admixture was not identified in Northern Europeans.[14]
  • Pino-Yanes et al. (2011) found that from an autosomal analysis, the average Northwest African influence is about 17% in Canary Islanders, with a wide interindividual variation ranging from 0% to 96%. The substantial Northwest African ancestry found for Canary Islanders supports the idea that, despite the aggressive conquest by the Spanish in the 15th century and the subsequent immigration, genetic footprints of the first settlers of the Canary Islands persist in the current inhabitants. Paralleling mtDNA findings, the largest average Northwest African contribution was found for the samples from La Gomera.[15]
  • Auton et al. (2009) found that South West Europe had the highest proportion in Europe of haplotypes that are shared with sub-Saharan Africa (represented by Yoruba), and significantly more relative to South East Europe.[16]

Sexual Chromosomes

Generally, markers and lineages used to characterize African admixture are those that are believed to be specific to Africa. There are also DNA polymorphisms that are shared between populations native to

mitochondrial haplogroup M1.[2]

With regard to the paternal haplogroup E1b1b and maternal haplogroup M1, derivatives of these clades have been observed in prehistoric human fossils excavated at the

Kelif el Boroud site near Rabat were found to carry the broadly-distributed paternal haplogroup T-M184 as well as the maternal haplogroups K1, T2 and X2, the latter of which were common mtDNA lineages in Neolithic Europe and Anatolia. These ancient individuals likewise bore the Berber-associated Maghrebi genomic component. This altogether indicates that the Late Neolithic Kelif el Boroud inhabitants were ancestral to contemporary populations in the area, but also likely experienced gene flow from Europe.[17]

Other lineages that are now found in Africa and Europe may have a common origin in Asia (e.g. Y haplogroups

T and U subclades). One subclade of haplogroup U, namely U6a1, is known to have expanded from northern and eastern Africa back into Europe[18][19] even though haplogroup U6 is considered to have originated in the Middle East. Other lineages are known to have moved from Europe directly into Africa, for example mitochondrial haplogroups H1 and H3.[20]
Such bidirectional migrations between Africa and Eurasia complicate the task of defining admixture.

Y-DNA

One proposed example of

E1b1b, which is thought to have emerged about 40,000 years ago in the Horn of Africa, and branches of it are thought to have migrated to the Middle East by 14,000 years ago during the late Pleistocene period.[21][10][22]

Entering the late

Neolithic transition. E1b1b1 lineages are found throughout Europe but are distributed along a south-to-north cline, with an E1b1b1a mode in the Balkans.[1][23][24][a][b]

In separate migrations, E lineages in the form of the E1b1b1b subclade appear to have entered Europe from Northwest Africa into Iberia. In a sample of European males, haplogroup E was observed at a frequency of 7.2%.

Sardinians, Sub-Saharan Y-DNA lineages A1b1b2b and E1a1 were found at a total of 1.0% (A1b1b2b 0.5% / E1a1 0.5%).[29]

Haplogroups A and B are thought to have been the predominant haplogroups in central and southern Africa prior to the Bantu Expansion. Currently these haplogroups are less common than E lineages. In a sample of 5,000 African men, haplogroup A had a frequency of 5%. Haplogroup A has rare occurrences in Europe, but recently the haplogroup was detected in seven indigenous British males with the same Yorkshire surname.[30]

E3b1 and its haplogroups E-M81 and E-M78 in North Africa, and E-M123 in the Near East

The subclade E3b1 (probably originating in northeastern Africa) has a wide distribution in North Africa, the Horn of Africa, the Middle East, and Europe. This haplogroup, in

Italian peninsula. Another subclade, E3b1b-M81 is associated with Berber populations and is commonly found in regions that have had historical gene flow with northern Africa, such as the Iberian Peninsula, the Canary Islands, and Sicily.[31]

North African Y-DNA E-M81 was found at a total of 41.1% among "pasiegos" from Cantabria, Spain. That is the highest frequency observed in Europe to date.[1] Estimates of Y-Chromosome ancestry vary. Using 1140 samples from throughout the Iberian peninsula, giving a proportion of 10.6% North African ancestry to the paternal composite of Iberians.[27][32] From an analysis of the Y-chromosome with 659 samples from Southern Portugal, 680 from Northern Spain, 37 samples from Andalusia, 915 samples from mainland Italy, and 93 samples from Sicily found significantly higher levels of North African male ancestry in Portugal, Spain and Sicily (7.1%, 7.7% and 7.5% respectively) than in peninsular Italy (1.7%).[33] Considering both some E-M78 subhaplogroups and the E-M81 haplogroup, the contribution of northern African lineages to the entire male gene pool of Iberia (barring Pasiegos), continental Italy, and Sicily can be estimated as 5.6%, 3.6%, and 6.6%, respectively.[34] Y-DNA lineages E-V12 and E-V22 have been associated with a Levantine source (represented by modern Lebanese), while North African haplogroup E-M81 shows an average frequency of 1.53% in the current Sicilian and Southern Italian genetic pool, but the typical Maghrebin core haplotype 13-14-30-24-9-11-13 has been found in only two out of the five E-M81 individuals.[35] These results, along with the negligible contribution from North-African populations revealed by the admixture-like plot analysis, suggest only a marginal impact of trans-Mediterranean gene flows on the current Sicilian and Southern Italian genetic pool.[35]

mtDNA

Iberia (Spain and Portugal), where frequencies as high as 22% have been reported, and some regions of Southern Italy, where frequencies as high as 2% and 3% have been found. About 65% of the European L lineages most likely arrived in rather recent historical times (Romanization period, Arab conquest of the Iberian Peninsula and Sicily, Atlantic slave trade) and about 35% of L mtDNAs form European-specific subclades, revealing that there was gene flow from Sub-Saharan Africa toward Europe as early as 11,000 years ago.[36]

Map (in the link) showing the distribution of Sub-Saharan mtDNA (shown in red) in Europe
Map is From Cerezo et al. 2012[36]
Universidad de Santiago de Compostela
Iberia (Spain & Portugal) having the highest amount and strongest concentration of Sub-Saharan mtDNA in Europe.

In

Muslim occupation and prehistoric migrations before the Muslim occupation would have been the source of these lineages. The highest frequencies of Sub-Saharan lineages found so far in Europe were observed in the comarca of Sayago (18.2%) which is "comparable to that described for the South of Portugal".[42]

In Italy, haplogroup L lineages are present at lower frequencies than in Iberia and are detected only in certain regions: Latium, Volterra,[43] Basilicata, and Sicily.[44]

In eastern Europe, haplogroup L lineages are present at very low frequencies. Though a high diversity of African mtDNA lineages have been detected, few lineages have accumulated enough mutations in Europe to form

Ashkenazi heritage from central and eastern Europe[46] and less frequently in non-Jewish Poles.[47] L2a lineages are widespread throughout Africa; as a result, the origins of this lineage are uncertain.[48]

Haplogroup M1 is also found in Europe at low frequencies. Haplogroup M1 had a frequency of 0.3%.[19]
The origins of haplogroup M1 have yet to be conclusively established.

A prehistoric episode is likely to be the main contributor to the sub-Saharan presence in Mediterranean Europe.[49]

Frequencies of haplogroup L lineages

Country Region Number tested Study %
Europe Continent-wide (excl. Tuscany) 10,589 Achilli et al. (2007)[43] 0.79%
South Iberia Spain & Portugal 310 Casas et al. (2006)[41] 7.40%
Spain Countrywide 312 Álvarez et al. (2007)[50] 2.90%
Spain Central Spain 50 Plaza et al. (2003)[51] 4.00%
Spain North-West Spain 216 Achilli et al. (2007)[43] 3.70%
Spain Basque Country 156 Achilli et al. (2007)[43] 0.64%
Spain Galicia 92 Pereira et al. (2005)[37] 3.30%
Spain Zamora 214 Álvarez et al. (2010)[42] 4.70%
Spain Sayago 33 Álvarez et al. (2010)[42] 18.18%
Spain Cordoba 108 Casas et al. (2006)[41] 8.30%
Spain Huelva 135 Hernandez et al. (2014) 5.70%
Spain Catalonia 101 Álvarez-Iglesias et al. (2009) 2.97%
Spain Balearic Islands 231 Picornell et al. (2005)[52] 2.20%
Spain Canary Islands 300 Brehm et al. (2003)[38] 6.60%
Portugal Countrywide 594 Achilli et al. (2007)[43] 6.90%
Portugal Countrywide 1429 Barral-Arca et al. (2016)[53] 6.16%
Portugal Countrywide 549 Pereira et al. (2005)[37] 5.83%
Portugal North 100 Pereira et al. (2010)[54] 5.00%
Portugal Center 82 Pereira et al. (2010)[54] 9.70%
Portugal Center 82 Plaza et al. (2003)[51] 6.10%
Portugal South 195 Brehm et al. (2003)[38] 11.30%
Portugal South 303 Achilli et al. (2007)[43] 10.80%
Portugal
Coruche
 160 Pereira et al. (2010)[54] 8.70%
Portugal Pias 75 Pereira et al. (2010)[54] 3.90%
Portugal Alcácer do Sal 50 Pereira et al. (2010)[54] 22.00%
Portugal Azores 179 Brehm et al. (2003)[38] 3.40%
Portugal Madeira 155 Brehm et al. (2003)[38] 12.90%
Portugal Madeira 153 Fernandes et al. (2006)[55] 12.40%
Italy Countrywide 583 Brisighelli et al. (2012)[31] 1.20%
Italy Countrywide 865 Boattini et al. (2013)[56] 0.00%
Italy Countrywide 240 Babalini et al. (2005)[57] 0.40%
Italy Tuscany 322 Achilli et al. (2007)[43] 1.86%
Italy Tuscany 49 Plaza et al. (2003)[51] 2.00%
Italy Volterra 114 Achilli et al. (2007)[43] 2.63%
Italy Latium 138 Achilli et al. (2007)[43] 2.90%
Italy Marche 813 Achilli et al. (2007)[43] 0.98%
Italy Central Italy 83 Plaza et al. (2003)[51] 1.20%
Italy Lombardy 177 Achilli et al. (2007)[43] 0.00%
Italy Piedmont 169 Achilli et al. (2007)[43] 0.00%
Italy Sardinia 258 Pardo et al. (2012)[58] 0.40%
Italy Sardinia 73 Plaza et al. (2003)[51] 2.80%
Italy Sardinia 85 Sanna et al. (2011)[59] 0.00%
Italy
Ogliastra
)		 475 Fraumene C et al. (2003)[60] 0.00%
Italy Sardinia 96 Morelli et al. (1999) 0.00%
Italy Campania (South Italy) 313 Achilli et al. (2007)[43] 0.32%
Italy Basilicata (South Italy) 92 Ottoni et al. (2009)[44] 2.20%
Italy Apulia & Calabria (South Italy) 226 Achilli et al. (2007)[43] 0.00%
Italy Southern Italy 115 Sarno et al. (2014)[35] 0.00%
Italy Southern Italy 37 Plaza et al. (2003)[51] 8.10%[failed verification]
Italy Sicily 106 Cali et al. (2003) 0.94%
Italy Sicily 105 Achilli et al. (2007)[43] 1.90%
Italy Sicily 169 Plaza et al. (2003)[51] 0.60%
Italy Sicily 198 Sarno et al. (2014)[35] 1.01%
Italy Sicily 465 Romano et al. (2003)[61] 0.65%
Greece Crete 202 Achilli et al. (2007)[43] 0.99%
Greece Crete 283 Martinez et al. (2008)[62] 0.00%
Greece Macedonia 125 Richards et al. (2000)[63] 0.00%
Greece Countrywide 155 Achilli et al. (2007)[43] 0.00%
Cyprus Cyprus 91 Irwin et al. (2008)[64] 3.30%[failed verification]
United Kingdom England 335 Achilli et al. (2007)[43] 0.60%
United Kingdom Wales 92 Achilli et al. (2007)[43] 0.00%
Finland Countrywide 121 Achilli et al. (2007)[43] 0.82%
Germany Countrywide 335 Achilli et al. (2007)[43] 0.30%
Ireland Countrywide 300 Achilli et al. (2007)[43] 0.00%
France Countrywide 332 Achilli et al. (2007)[43] 0.30%
Bulgaria Countrywide 141 Achilli et al. (2007)[43] 0.71%
Bosnia and Herzegovina Countrywide 144 Achilli et al. (2007)[43] 0.69%

In an analysis which also contains an admixture data but no cluster membership coefficients, shows little to no Sub-Saharan African influence in a wide array of European samples, i.e. Albanians, Austrians, Belgians, Bosnians, Bulgarians, Croatians, Cypriots, Czechs, Danes, Finns, Frenchmen, Germans, Greeks, Hungarians, Irish, Italians, Kosovars, Lithuanians, Latvians, Macedonians, Netherlanders, Norwegians, Poles, Portuguese, Romanians, Russians, Scots, Serbians, Slovaks, Slovenians, Spaniards, Swedes, Swiss (German, French and Italian), Ukrainians, subjects of the United Kingdom, and Yugoslavians.[16]

U6 is most common in Spain and Portugal.[66][51]

Frequencies of haplogroup U6 lineages

Country Region Number tested Study %
Italy Countrywide 583 Brisighelli et al. (2012)[31] 0.8%
Italy Mainland 411 Plaza et al. (2003)[51] 0.0%
Italy Countrywide 865 Boattini et al. (2013)[56] 0.35%
Italy Sicily 169 Plaza et al. (2003)[51] 0.6%
Italy Sicily 106 Maca-Meyer et al. (2003).[66] 0.94%
Italy Lazio 52 Babalini et al. (2005)[57] 5.8%
Italy Abruzzo (Molise) 73 Babalini et al. (2005)[57] 0%
Italy Campania 48 Babalini et al. (2005)[57] 0%
Italy Volterra (Tuscany) 114 Achilli et al. (2007)[43] 0.00%
Italy Murlo (Tuscany) 86 Achilli et al. (2007)[43] 1.20%
Italy Casentino (Tuscany) 122 Achilli et al. (2007)[43] 0.80%
Italy Sicily 105 Achilli et al. (2007)[43] 0.95%
Italy Latium 138 Achilli et al. (2007)[43] 0.00%
Italy Lombardy 177 Achilli et al. (2007)[43] 0.00%
Italy Piedmont 169 Achilli et al. (2007)[43] 0.00%
Italy Marche 813 Achilli et al. (2007)[43] 0.25%
Italy Campania 313 Achilli et al. (2007)[43] 1.28%
Italy Apulia-Calabria 226 Achilli et al. (2007)[43] 1.33%
Italy Sardinia 370 Achilli et al. (2007)[43] 0.27%
Spain Central Spain 50 Plaza et al. (2003)[51] 2.0%
Spain Galicia 103 Plaza et al. (2003)[51] 1.9%
Spain Galicia 135 Maca-Meyer et al. (2003)[66] 2.2%
Spain Catalonia 118 Maca-Meyer et al. (2003)[66] 1.6%
Spain Huelva 135 Hernandez et al. (2014)[67] 8.8%
Spain Maragatos 49 Maca-Meyer et al. (2003)[66] 8.1%
Spain Canary Islands 300 Brehm et al. (2003)[38] 14.0%
Portugal Countrywide 54 Plaza et al. (2003)[51] 5.6%
Portugal North Portugal 184 Maca-Meyer et al. (2003)[66] 4.3%
Portugal Central Portugal 161 Brehm et al. (2003)[38] 1.9%
Portugal Madeira 155 Brehm et al. (2003)[38] 3.9%
Portugal Madeira 153 Fernandes et al. (2006)[55] 3.3%
Iberia Spain & Portugal 887 Plaza et al. (2003)[51] 1.8%

GM immunoglobulin allotypes

Further studies have shown that the presence of haplotype GM*1,17 23' 5* in southern Europe. This haplotype is considered a genetic marker of Sub-Saharan Africa, where it shows frequencies of about 80%.

Aran valley in the Pyrenees.[70] Although some researchers have associated African traces in Iberia to Islamic conquest, the presence of GM*1,17 23' 5* haplotype in Iberia may in fact be due to more ancient processes as well as more recent ones through the introduction of genes from slaves sold from Africa.[69]

In Sicily the North African haplotype Gm 5*;1;17; ranges from 1.56% at Valledolmo to 5.5% at Alia.[71] The hypothesis is that the presence of this haplotype suggests past contacts with people from North Africa. The introduction of African markers could be due to the Phoenician colonization at the end of the second millennium B.C. or to the more recent Arab conquest (8th–9th centuries A.D.).

Paleoanthropology

The migration of farmers from the Middle East into Europe is believed to have significantly influenced the genetic profile of present-day Europeans. Some recent studies have focused on corroborating current genetic data with the archeological evidence from Europe, the Middle East, and Africa.[25] The Natufian culture, which existed about 12,000 years ago, has been the subject of various archeological investigations, as it is generally believed to be the source of the European and North African Neolithic.

According to one hypothesis,

Kebaran, a culture indigenous to the Levant, and (2) the Mushabian, a culture introduced into the Levant from North Africa†. It is suggested that the Mushabian culture originated in Africa, given that archeological sites with Mushabian industries in the Nile Valley predate those in the Levant†. The Mushabians would have then moved into the Sinai from the Nile Delta bringing with them their technologies†. The overpopulation in Northeast Africa contributed to the development of the Natufian adaptation, which resulted in agriculture becoming a new way of sustenance.[9]

From an analysis of human remains from the Natufian culture, there is evidence of Sub-Saharan influences in the Natufian samples.

E1b1b lineages from Northeast Africa to the Levant and then into Europe.[72]

According to an ancient DNA analyse on Natufian skeletal remains from present-day northern Israel, the Natufians in fact shared no evident genetic affinity to sub-Saharan Africans.

hunter gatherers. Mating between Natufians, other Neolithic Levantines, Caucasus Hunter Gatherers (CHG), Anatolian and Iranian farmers is believed to have decreased genetic variability
among later populations in the Middle East. The scientists suggest that the Levantine early farmers may have spread southward into East Africa, bringing along Western Eurasian and Basal Eurasian ancestral components separate from that which would arrive later in North Africa.

† The Mushabian industry is now known to have originated in the Levant from the previous lithic industries of the region of Lake Lisan.[79] The Mushabian industry was originally thought to have originated in Africa because the microburin technique was not yet known to be much older in the eastern Levant.[80] Currently there is no known industry to connect with the African migration that occurred 14,700 years ago,[1] but it no doubt caused a population expansion in the Negev and Sinai which would not have accommodated an increase in population with the meager resources of a steppe/desert climate.[9] Since all of the known cultures in the Levant at the time of the migration originated in the Levant and an archaeological culture cannot be associated with it, there must have been assimilation into a Levantine culture at the onset, most likely the Ramonian which was present in the Sinai 14,700 years ago.[81]

See also

Notes

  1. ^ Recently, it has been proposed that E3b originated in eastern Africa and expanded into the Near East and northern Africa at the end of the Pleistocene. E3b lineages would have then been introduced from the Near East into southern Europe by migrant farmers, during the Neolithic expansion.[1]
  2. ^ A Mesolithic population carrying Group III lineages with the M35/M215 mutation expanded northwards from sub-Saharan to north Africa and the Levant. The Levantine population of farmers that dispersed into Europe during and after the Neolithic carried these African Group III M35/M215 lineages, together with a cluster of Group VI lineages characterized by M172 and M201 mutations.[24]

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    PMID 14563219
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  67. PMID 24460736
    .
  68. S2CID 38748780
    .
  69. ^
    PMID 17597520
    .
  70. PMID 11851984
    .
  71. S2CID 9663858
    .
  72. S2CID 25142338
    .
  73. ^ Lazaridis 2016 Quote: "However, no affinity of Natufians to sub-Saharan Africans is evident in our genome-wide analysis, as present-day sub-Saharan Africans do not share more alleles with Natufians than with other ancient Eurasians (Extended Data Table 1)."
  74. PMID 27459054
    ., Supplementary Table 1.
  75. PMID 27459054
    .
  76. PMID 27417496
    .
  77. PMID 27502179
    .
  78. PMID 24901650
    .
  79. ISBN 978-1-84217-340-4
    .
  80. doi:10.3406/paleo.2006.5168
    .
  81. S2CID 162887983
    .

Further reading

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