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Home , Acheulean, Ancient North Eurasian, Aurignacian, Gravettian, Hand axe, Magdalenian

Outline

Early Modern I and Alan R. Rogers I Expansion out of I

I Eurasia February 7, 2018

1 / 71 2 / 71 Moderns invade Eurasia

I

I 2.6–1.7 mya

I Earliest stone tools

I Probably made by habilis/rudolphensis

3 / 71 4 / 71 (1.5–0.3 mya)

Oldowan tools 2.6–1.7 mya

5 / 71 6 / 71 Acheulean tools (1.5–0.3 mya) Distribution of Acheulean tools

7 / 71 8 / 71 Acheulean tools tools

I Associated with /erectus

I 1.5–0.3 myr ago I 300–30 kya

I Essentially unchanged for 1.2 myr I and I Wide distribution in space western I Neandertals

9 / 71 10 / 71 Neandertal sites Mousterian tools

I Neandertals

I 300–30 kya

I Little change in 270 kyr

I Wide distribution in space

11 / 71 12 / 71 tools

I 19–15 kya

Aurignacian tools I Laurel-leaf I 34–23 kya points

I Earliest Upper I Later , Europe Paleolithic, I Modern humans Europe I Modern humans

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LASCAUX 17 kya

Grotte Chauvet 31 kya

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Art Venus of Deer’ head Lespugue 18–10 kya (28 kya)

17 / 71 18 / 71 thrower

I increases force of throw INITIAL I penetrated Spanish armor UPPER PALE- I kill at 70 m OLITHIC I by 14 kya OF EUROPE

19 / 71 20 / 71 traditions of the Upper Paleolithic

Neandertal & Modern Modern (at right):

Age Tradition I small 35–29 kya Chatelperronian I Change is rapid. I long-limbed 34–23 kya I Each tradition occupies a I gracile 28–22 kya small region. 19–15 kya Solutrean I flat face I varies in space. 18–10 kya I chin

I forehead

I domed skull

21 / 71 22 / 71 Herto, (160 kya) Manot, (55 kya)

Modern

But with archaic features, such as occipital bun.

23 / 71 24 / 71 Outline Comas et al (1997) studied European mismatch distributions

Archaeology and paleontology ◦ I Expansion out of Africa

I Paleolithic Eurasia

I Mesolithic Eurasia

25 / 71 26 / 71 Mismatch distributions suggest expansion across Europe Nuclear DNA: Xing et al 2010

Mid-East and Turkey: early expansions. I Studied large sample of nuclear DNA from populations in British, : late expansions Africa, Europe, East Asia, and India. I Eurasian populations left Africa about 100 thousand ago Paleolithic or ? I Did not separate until 25–40 thousand years ago Comas et al thought Paleolithic but may have been misled by mtDNA clock.

27 / 71 28 / 71 Outline

Archaeology and paleontology ◦ Expansion out of Africa ◦ I Paleolithic Eurasia

I Mesolithic Eurasia

29 / 71 30 / 71 Paleolithic DNA of NW Eurasia

Ust’-Ishim: a 45-ky-old modern man from W

Excavated 2008

Fu et al. (2014)

Note time scale at back.

31 / 71 32 / 71 Location of Ust’-Ishim Dates and temperature

1, Ust’-Ishim; 4, Denisova.

33 / 71 34 / 71 History of population size

Ust’-Ishim PC map

Ust’-Ishim (purple) similar to Central Asians

Red, assumes Ust’-Ishim lived today; Blue, fits curve to others and estimates mutation rate. Lived during late bottleneck.

35 / 71 36 / 71 allele sharing

Conventional wisdom Problem

I Early emigration out of Ust’-Ishim is equally related to Africa to Australia & New northern and southern modern Guinea. populations.

I Later northern emigration. Perhaps it was a 3-way split, and I Ust’-Ishim should be part of Ust’-Ishim’s population went northern population. extinct.

Ust’-Ishim has 2.3% Neanderthal DNA—in big chunks. Implies recent admixture (50–60 kya).

37 / 71 38 / 71 Man from Pe¸stera cu Oase,

40 ky-old Oase 1 6–9% of genome is Neanderthal—3 larger than any × other

3 Neanderthal segments >50 cM

Implies Neanderthal ancestor 4–6 generations back.

Shares more derived alleles with E Asians & Native Americans than with Europeans.

Fu et al. (2015) Green: Oase alleles shared with Neanderthal.

39 / 71 40 / 71 Kostenki 14: a 37 ky old Russian man Affinities of Kostenki with world populations

(Seguin-Orlando et al. 2014) Similar to Northern Europeans.

41 / 71 42 / 71 Affinities of Kostenki with European populations Neandertal allele sharing

(Seguin-Orlando et al. 2014)

(Seguin-Orlando et al. 2014) K14 has more Neandertal DNA and in bigger chunks. Similar to northern and western Europeans.

43 / 71 44 / 71 Neandertal admixture fraction 24 ky old burial from Mal’ta, Siberia

Kostenki (K14) had more Neanderthal DNA than modern humans and in bigger chunks. Implies recent admixture–54 kya.

(Seguin-Orlando et al. 2014)

(Raghavan et al. 2013)

45 / 71 46 / 71 Location of Mal’ta site Mal’ta in principal components map

Raghavan et al. (2013)

47 / 71 48 / 71 (Raghavan et al. 2013) Affinities of Mal’ta with other populations 24 ky old burial from Mal’ta, Siberia

I 1/3 of ancestry shared with Native Americans and Europeans

I European mitochondrial DNA (Raghavan et al. 2013)

(Raghavan et al. 2013) Similar to Amerindians and Northern Europeans.

49 / 71 50 / 71 mtDNA haplogroups: 45–25 kya mtDNA haplogroups: 19.5–14.5 kya

(Posth et al 2016) (Posth et al 2016)

51 / 71 52 / 71 mtDNA haplogroups: 14.5–11.5 kya mtDNA haplogroups: 11.5–7 kya

(Posth et al 2016) (Posth et al 2016)

53 / 71 54 / 71 Outline

Two big changes in variation.

One coincides with Archaeology and paleontology last glacial ◦ Expansion out of Africa maximum (LGM). ◦ Paleolithic Eurasia ◦ The other with the I Mesolithic Eurasia last glacial (aka the ).

(Posth et al 2016)

55 / 71 56 / 71 La Bra˜na:a 7,000 y old forager La Bra˜nasite

57 / 71 58 / 71 La Bra˜naPC Map: World La Bra˜naPC Map: Europe

On world-wide But La Bra˜nais not scale, La Bra˜na is like modern European. Europeans.

59 / 71 60 / 71 Neandertal admixture fraction Gradual decline in Neanderthal admixture

La Bra˜nahad more Neanderthal DNA than modern humans.

(Seguin-Orlando et al. 2014)

61 / 71 62 / 71 La Bra˜naadaptations Study of Lazaridis et al (2014)

I Dark skin

I Blue eyes DNA sequences of: Label Location Samples Age Description I Lactose intolerant Loschbour Luxembourg 1 8 ky forager I Poor at digesting starch. Motala Sweden 7 8 ky forager I Modern allele at 24 of 40 loci affecting immune function. Stuttgart Germany 1 7 ky farmer These observations suggest that the 2345 0 ky modern transition did not drive all cases of adaptive The moderns were not sequenced but were genotyped at 594,924 innovation on immunity genes found in modern autosomal single-nucleotide polymorphisms. Europeans. (Olalde et al 2014)

63 / 71 64 / 71 Lazaridis et al samples Population size

Loschbour, Motala: 8 ky old foragers; Stuttgart: 7 ky old farmer. Loschbour: 8 ky old forager; Stuttgart: 8 kya farmer; others: modern. No post-Pleistocene growth of forager population.

65 / 71 66 / 71 Contributions to modern Europeans PCA map

ANE WHG West-European hunter-gatherer EEF Early European Farmer

67 / 71 68 / 71 Contributions to modern Europeans Adaptations

I Lochbour (forager) relatively good at digesting starch, even before agriculture.

I Stuttgart (farmer): Dark hair, , brown eyes.

I Foragers: Dark skin, hair and blue eyes.

I But one Motala (forager) sample had one copy of light skin allele. It was in Europe before agriculture.

69 / 71 70 / 71 Summary

I Archaics changed slowly both in time and space. Pace of change is faster for modern humans.

I Moderns appear first in Africa, 160 ky ago. Spread through Eurasia 50 ky ago. Introduce , tools, projectile ∼ weapons.

I Genome of Ust’-Ishim, a 45-ky-old Siberian, was modern, distinct from Africans, but no closer to northern than to southern non-Africans. Big chunks of Neanderthal genome imply recent admixture.

I Kostenki, a 36 ky-old Siberian, genetically similar to Europeans. More Neanderthal DNA than modern humans.

I Mal’ta

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