Hominid dispersals in Europe

Short description: Colonisation of the European continent

Hominid dispersals in Europe refers to the colonisation of the European continent by various species of hominid, including hominins and archaic and modern humans. Prior to arriving in Europe, the source of hominids appeared to be East Africa, where stone tools and hominid fossils are the most abundant and recorded.[1]

Short and repetitive migrations of archaic humans before 1 million years ago suggest that their residence in Europe was not permanent at the time.[2] Colonisation of Europe in prehistory was not achieved in one immigrating wave, but instead through multiple dispersal events.[3] Most of these instances in Eurasia were limited to 40th parallel north.[3] Besides the findings from East Anglia, the first constant presence of archaic humans in Europe begins 500,000–600,000 years ago.[4] However, this presence was limited to western Europe, not reaching places like the Russian plains, until 200,000–300,000 years ago.[4] The exception to this was discovered in East Anglia, where archaic humans briefly inhabited 700,000 years ago.[5]

Arising in Europe at least 400,000 years ago, the Neanderthal archaic humans (a descendant of Homo heidelbergensis) would become more stable residents of the continent. The Cro-Magnon early modern humans would arrive about 50,000 years ago, leading to the extinction of the Neanderthals about 37,000 years ago, following interbreeding between the two groups.

## Contents

* 1 Pre-human hominids
* 2 Archaic humans
* 3 Anatomically modern humans
* 4 Pressures favoring migration
* 5 See also
* 6 References
* 7 External links

## Pre-human hominids

A fragment of skull belonging to Ouranopithecus macedoniensis, a hominid found in Europe in the Late Miocene[6]

Short description: Hominin events for the last 10 million years

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(million years ago)

(See also: Life timeline, and Nature timeline.)

}} In the Early Miocene, Europe had a subtropical climate and was intermittently connected to Africa by land bridges. At the same time, Africa was becoming more arid, prompting the dispersal of its tropical fauna—including primates—north into Europe.[7] Apes first appear in the European fossil record 17 million years ago with Griphopithecus.[8] The closely related Kenyapithecus is also known from fossils in Germany, Slovakia and Turkey.[7] Both Griphopithecus and Kenyapithecus are considered likely to be ancestral to the great apes.[9] From 13 million to 9 million years ago, hominids flourished in Europe and underwent an adaptive radiation as they diversified in response to a gradually cooling climate.[7][10] Middle Miocene European hominids include Pierolapithecus, Anoiapithecus, Dryopithecus, Hispanopithecus, and Rudapithecus.[6] The diversity and early appearance of great apes in Europe has led some scientists to theorise that hominids in fact evolved there, before dispersing "back to Africa" in the Middle Miocene.[7][9]

Around 9 million years ago most of Europe's hominid species fell victim to the Vallesian crisis, an extinction event caused by the disappearance of the continent's forests.[7][10] Some hominid species survived the event: Oreopithecus, which became isolated in forest refugia; and Ouranopithecus, which adapted to the open environments of the late Miocene.[7] However, both were extinct by 7 million years ago.[6]

In 2017, a reanalysis of Graecopithecus fossils from Greece and Bulgaria, previously associated with Ouranopithecus, concluded that the species was in fact a hominin dating to just after the last common ancestor of humans and chimpanzees (about 7.2 million years ago).[11] The authors suggested that the origins of the human lineage were therefore in the Mediterranean, not Africa.[12][13][14] Others are sceptical of their claims.[14][15][16]

Although subtropical conditions returned to Europe in the Pliocene (5.33–2.58 million years ago), there are no known fossil hominids from this period.[17]

## Archaic humans

Further information: Dmanisi skull 3

Homo erectus populations lived in southeastern Europe by 1.8 million years ago.[18]

The most archaic human fossils from the Middle Pleistocene (780,000–125,000 years ago)[19] have been found in Europe. Remains of Homo heidelbergensis have been found as far north as the Atapuerca Mountains in Gran Dolina, Spain, and the oldest specimens can be dated from 850,000 to 200,000 years ago.[20][21]

Neanderthals evolved from a branch of Homo heidelbergensis that migrated to Europe during the Middle Pleistocene.[22] Neanderthal populations date back at least as far as 400,000 years ago in the Atapuerca Mountains, Spain.[23] While lacking the robustness attributed to west European Neanderthal morphology, other populations did inhabit parts of eastern Europe and western Asia.[23] Between 45,000–35,000 years ago, modern humans (Homo sapiens) replaced all Neanderthal populations in Europe anatomically and genetically.[24] This is evident in the transfer and combination of technology and culture.

## Anatomically modern humans

Further information: Genetic history of Europe

The recent expansion of anatomically modern humans reached Europe around 40,000 years ago, from Central Asia and the Middle East, as a result of cultural adaption to big game hunting of sub-glacial steppe fauna.[25][26] Neanderthals were present both in the Middle East and in Europe, and the arriving populations of anatomically modern humans (also known as "Cro-Magnon" or European early modern humans) have interbred with Neanderthal populations to a limited degree.

Modern human remains dating to 43,000–45,000 years ago have been discovered in Italy[27] and Britain,[28] with the remains found of those that reached the European Russian Arctic 40,000 years ago.[29][30] The composition of European populations was later altered by further migrations, notably the Neolithic expansion from the Middle East, and still later the Chalcolithic population movements associated with Indo-European expansion.

The dominant population of Europe is composed of three major foundational populations, dubbed "Western Hunter-Gatherers" (WHG), "Early European Farmers" (EEF) and "Ancient North Eurasian" (ANE). WHG represents the remnant of the original Cro-Magnon population after they re-peopled Europe after the Last Glacial Maximum. EEF represents the introgression of Near Eastern populations during the Neolithic Revolution, and ANE is associated both with the Mesolithic Uralic expansion to Northern Europe and the Indo-European expansion to Europe in the Chalcolithic.[31]

## Pressures favoring migration

Homo ergaster specimens indicate a change toward a diet more reliant on animal products, evident by greater encephalization with higher energy requirements.[32] This transition to becoming more carnivorous affected the way of life unlike primates before.[33][34] Archaeological evidence of cut bones from large mammals and broken stone tools increasing in frequency support this increasing trend.[4] To meet increasing demand of calories, the range of hominids would have expanded, making the necessary hunting versus prior scavenging possible.[4] It is believed that the adjustments required to meet these new demands would expand the home range size eight to ten times.[35] Range could also increase or decrease in size due to environmental changes.[4] A more recent example is absence of humans in Britain during the last glacial maximum which ended in the Late Pleistocene, 10,000 years ago.[36][37] At this time, Russia had an influx of people following the major prey species shifting to this region.[38] It has been argued that Neanderthals', and previous hominids', expansion northward were limited by lacking proper thermoregulation.[4] Behavioural adaptations such as clothes-making to overcome the cold is evident in archaeological finds.[4] The potential to expand also grew with the Neanderthal reaching the status of top carnivores.[4] These humans could fear less during expansion, without the worry of other predators. The desire to push into these northern areas was influenced by this requirement to eat a lot of meat to satisfy the human brain which uses 20% of the body's energy.[39] Larger game for hunting is available the closer you are to the poles.[4]

## See also

* Early human migrations
* Historical migration

* History of Europe
* Human migration

## References

1. ↑ Roebroeks, Wil (2006). "The human colonisation of Europe: where are we?". Journal of Quaternary Science 21 (5): 425–435. doi:10.1002/jqs.1044. Bibcode: 2006JQS....21..425R.
2. ↑ Anton, S.C.; Swisher C. (2004). "Early dispersals of Homo from Africa.". Annual Review of Anthropology 33: 271–296. doi:10.1146/annurev.anthro.33.070203.144024.
3. ↑ 3.0 3.1 Dennell, R.W. (2003). "Dispersal and colonisation, long and short chronologies: how continuous is the Early Pleistocene record for hominids outside East Africa?". Journal of Human Evolution 45 (6): 421–440. doi:10.1016/j.jhevol.2003.09.006. PMID 14643672. Bibcode: 2003JHumE..45..421D.
4. ↑ 4.0 4.1 4.2 4.3 4.4 4.5 4.6 4.7 4.8 Roebroeks, Wil (2006). "The human colonisation of Europe: where are we?". Journal of Quaternary Science 21 (5): 425–435. doi:10.1002/jqs.1044. Bibcode: 2006JQS....21..425R.
5. ↑ Henderson, Mark (June 4, 2002). "'Anglia Man' becomes earliest Ancient Briton". The Times: British News. http://www.archaeology.ws/2002-6-6.htm.
6. ↑ 6.0 6.1 6.2 Harrison, Terry (2016). "Miocene Primates". in Fuentes, Augustin (in en). The International Encyclopedia of Primatology. John Wiley & Sons, Inc.. pp. 1–5. doi:10.1002/9781119179313.wbprim0227. ISBN 978-1-119-17931-3.
7. ↑ 7.0 7.1 7.2 7.3 7.4 7.5 Begun, David R.; Nargolwalla, Mariam C.; Kordos, László (2012). "European Miocene Hominids and the Origin of the African Ape and Human Clade" (in en). Evolutionary Anthropology: Issues, News, and Reviews 21 (1): 10–23. doi:10.1002/evan.20329. ISSN 1520-6505. PMID 22307721.
8. ↑ Heizmann, Elmar P.J.; Begun, David R. (2001). "The oldest Eurasian hominoid". Journal of Human Evolution 41 (5): 463–481. doi:10.1006/jhev.2001.0495. PMID 11681862. Bibcode: 2001JHumE..41..463H. http://doc.rero.ch/record/13528/files/PAL_E351.pdf.
9. ↑ 9.0 9.1 Moyà-Solà, Salvador; Alba, David M.; Almécija, Sergio; Casanovas-Vilar, Isaac; Köhler, Meike; Esteban-Trivigno, Soledad De; Robles, Josep M.; Galindo, Jordi et al. (2009). "A unique Middle Miocene European hominoid and the origins of the great ape and human clade" (in en). Proceedings of the National Academy of Sciences 106 (24): 9601–9606. doi:10.1073/pnas.0811730106. ISSN 0027-8424. PMID 19487676. Bibcode: 2009PNAS..106.9601M.
10. ↑ 10.0 10.1 DeMiguel, Daniel; Alba, David M.; Moyà-Solà, Salvador (2014). "Dietary Specialization during the Evolution of Western Eurasian Hominoids and the Extinction of European Great Apes". PLOS ONE 9 (5). doi:10.1371/journal.pone.0097442. ISSN 1932-6203. PMID 24848272. Bibcode: 2014PLoSO...997442D.
11. ↑ Fuss, Jochen; Spassov, Nikolai; Begun, David R.; Böhme, Madelaine (2017). "Potential hominin affinities of Graecopithecus from the Late Miocene of Europe". PLOS ONE 12 (5). doi:10.1371/journal.pone.0177127. ISSN 1932-6203. PMID 28531170. Bibcode: 2017PLoSO..1277127F.
12. ↑ "7.2-million-year-old pre-human remains found in the Balkans: New hypothesis about the origin of humankind suggests oldest hominin lived in Europe" (in en). ScienceDaily. https://www.sciencedaily.com/releases/2017/05/170523083548.htm.
13. ↑ "Graecopithecus freybergi: Oldest Hominin Lived in Europe, not Africa | Paleoanthropology | Sci-News.com" (in en-US). Breaking Science News | Sci-News.com. http://www.sci-news.com/othersciences/anthropology/graecopithecus-freybergi-hominin-04888.html.
14. ↑ 14.0 14.1 "Our common ancestor with chimps may be from Europe, not Africa" (in en-US). New Scientist. https://www.newscientist.com/article/2132026-our-common-ancestor-with-chimps-may-be-from-europe-not-africa/.
15. ↑ Benoit, Julien. "There's not enough evidence to back the claim that humans originated in Europe" (in en). The Conversation. http://theconversation.com/theres-not-enough-evidence-to-back-the-claim-that-humans-originated-in-europe-78280.
16. ↑ Curnoe, Darren (May 23, 2017). "Did humans evolve in Europe rather than Africa? We don't have the answer just yet". Phys.org. https://phys.org/news/2017-05-humans-evolve-europe-africa-dont.html.
17. ↑ Eronen, Jussi T.; Rook, Lorenzo (2004). "The Mio-Pliocene European primate fossil record: dynamics and habitat tracking". Journal of Human Evolution 47 (5): 323–341. doi:10.1016/j.jhevol.2004.08.003. PMID 15530351. Bibcode: 2004JHumE..47..323E. http://doc.rero.ch/record/15618/files/PAL_E3067.pdf.
18. ↑ Lewis, Barry; Jurmain, Robert; Kilgore, Lynn (2012). "10". in Mark Kerr. Understanding Humans: An Introduction to Physical Anthropology and Archaeology. Belmont, California: Wadsworth. pp. 233–234. ISBN 978-1-111-83177-6.
19. ↑ Lewis, Barry; Jurmain, Robert; Kilgore, Lynn (2012). "11". in Mark Kerr. Understanding Humans: An Introduction to Physical Anthropology and Archaeology. Belmont, California: Wadsworth. p. 252. ISBN 978-1-111-83177-6.
20. ↑ Rightmire, G.P. (7 December 1998). "Human evolution in the Middle Pleistocene: The role of Homo heidelbergensis". Evolutionary Anthropology 6 (6): 218–227. doi:10.1002/(sici)1520-6505(1998)6:6<218::aid-evan4>3.0.co;2-6.
21. ↑ Lewis, Barry; Jurmain, Robert; Kilgore, Lynn (2012). "11". in Mark Kerr. Understanding Humans: An Introduction to Physical Anthropology and Archaeology. Belmont, California: Wadsworth. pp. 254–257. ISBN 978-1-111-83177-6.
22. ↑ Lewis, Barry; Jurmain, Robert; Kilgore, Lynn (2012). "11". in Mark Kerr. Understanding Humans: An Introduction to Physical Anthropology and Archaeology. Belmont, California: Wadsworth. pp. 256–257. ISBN 978-1-111-83177-6.
23. ↑ 23.0 23.1 Lewis, Barry; Jurmain, Robert; Kilgore, Lynn (2012). "11". in Mark Kerr. Understanding Humans: An Introduction to Physical Anthropology and Archaeology. Belmont, California: Wadsworth. pp. 262–269. ISBN 978-1-111-83177-6.
24. ↑ Mellars, Paul; French, Jennifer C. (29 July 2011). "Tenfold Population Increase in Western Europe at the Neandertal-to-Modern-Human Transition". Science 333 (6042): 623–627. doi:10.1126/science.1206930. PMID 21798948. Bibcode: 2011Sci...333..623M.
25. ↑ Oppenheimer, Stephen "Out of Eden: Peopling of the World" (Robinson; New Ed edition (March 1, 2012))
26. ↑ Hoffecker, J. (2006). A Prehistory of the North: Human Settlements of the Higher Latitudes. New Jersey: Rutgers University Press. p. 101. https://archive.org/details/prehistorynorthh00hoff. [ISBN missing]
27. ↑ Benazzi, S.; Douka, K.; Fornai, C.; Bauer, C.C.; Kullmer, O.; Svoboda, J.Í.; Pap, I.; Mallegni, F. et al. (2011). "Early dispersal of modern humans in Europe and implications for Neanderthal behaviour". Nature 479 (7374): 525–8. doi:10.1038/nature10617. PMID 22048311. Bibcode: 2011Natur.479..525B.
28. ↑ Higham, T.; Compton, T.; Stringer, C.; Jacobi, R.; Shapiro, B.; Trinkaus, E.; Chandler, B.; Gröning, F. et al. (2011). "The earliest evidence for anatomically modern humans in northwestern Europe". Nature 479 (7374): 521–4. doi:10.1038/nature10484. PMID 22048314. Bibcode: 2011Natur.479..521H.
29. ↑ Pavlov, Pavel; Svendsen, John Inge; Indrelid, Svein (2001). "Human presence in the European Arctic nearly 40,000 years ago". Nature 413 (6851): 64–7. doi:10.1038/35092552. PMID 11544525. Bibcode: 2001Natur.413...64P.
30. ↑ "Mamontovaya Kurya:an enigmatic, nearly 40000 years old Paleolithic site in the Russian Arctic". https://notendur.hi.is/oi/AG-326 2006 readings/Russian Arctic/Svendsen_MAMMOTH2003.pdf.
31. ↑ Lazaridis, Iosif (2014). "Ancient human genomes suggest three ancestral populations for present-day Europeans". Nature 513 (7518): 409–413. doi:10.1038/nature13673. PMID 25230663. Bibcode: 2014Natur.513..409L. . "most present Europeans derive from at least three highly differentiated populations: West European Hunter-Gatherers (WHG), who contributed ancestry to all Europeans but not to Near Easterners; Ancient North Eurasians (ANE) related to Upper Paleolithic Siberians, who contributed to both Europeans and Near Easterners; and Early European Farmers (EEF), who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model these populations' deep relationships and show that EEF had ~44% ancestry from a "Basal Eurasian" population that split prior to the diversification of other non-African lineages."
32. ↑ Aiello, L.C.; Wheeler, P.E. (1995). "The Expensive Tissue Hypothesis: the brain and the digestive system in human and primate evolution.". Current Anthropology 36 (2): 199–221. doi:10.1086/204350.
33. ↑ Stanford, C.B. (March 1996). "The hunting ecology of wild chimpanzees: implications for the evolutionary ecology of Pliocene hominids.". American Anthropologist 98 (1): 96–113. doi:10.1525/aa.1996.98.1.02a00090.
34. ↑ Stanford, C. (2001). Bunn, H. ed. "A comparison of social meat-foraging by chimpanzees and human foragers". InMeat-eating and Human Evolution: 122–140. doi:10.1093/oso/9780195131390.003.0007. ISBN 978-0-19-513139-0.
35. ↑ Anton, S.C.; Leonard W.R.; Robertson M.L. (2002). "An ecomorphological model of the initial hominid dispersal from Africa.". Journal of Human Evolution 43 (6): 773–785. doi:10.1006/jhev.2002.0602. PMID 12473483. Bibcode: 2002JHumE..43..773A.
36. ↑ Terberger, T; Street, M. (2002). "Hiatus or continuity? New results for the question of pleniglacial settlement in Central Europe.". Antiquity 76 (293): 691–698. doi:10.1017/s0003598x00091134.
37. ↑ Roebroeks, W.; Speleers, B. (2002). Tuffreau, A. ed. "Last interglacial (Eemian) occupation of the North European plain and adjacent areas.". In le Dernier Interglaciaire et les Occupations Humaines du Paléolithique Moyen: 31–39.
38. ↑ Stuart, A.J.; Kosintsev P.A.; Higham T.F.G.; Lister A.M. (2004). "Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth.". Nature 431 (7009): 684–689. doi:10.1038/nature02890. PMID 15470427. Bibcode: 2004Natur.431..684S. http://doc.rero.ch/record/13494/files/PAL_E274.pdf.
39. ↑ Sorensen, M.V.; Leonard, W.R. (2001). "Neandertal energetics and foraging efficiency.". Journal of Human Evolution 40 (6): 483–495. doi:10.1006/jhev.2001.0472. PMID 11371151. Bibcode: 2001JHumE..40..483S.

## External links

* The Settlement and Colonization of Europe

* v
* t
* e

Early human migrations

Hominin and Homo dispersals

Lower Paleolithic|

* Out of Africa I
* Neanderthal range
* Europe
* Southeast Asia (Burma
* Philippines)

Middle Paleolithic|

* Africa
* Out of Africa II
* Coastal migration
* archaic admixture
* Australia
* Burma
* Philippines
* Europe

Upper Paleolithic|

* LGM refugia
* Mammoth steppe
* Ancient Beringian
* Settlement of the Americas
* Ancient North Eurasians
* Western hunter-gatherers
* Younger Dryas
* Burma
* Philippines

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