Outline Early Modern Humans I Archaeology and paleontology Alan R. Rogers I Expansion out of Africa I Paleolithic Eurasia I Mesolithic Eurasia February 7, 2018 1 / 71 2 / 71 Moderns invade Eurasia I Oldowan tools I 2.6–1.7 mya I Earliest stone tools I Probably made by Homo habilis/rudolphensis 3 / 71 4 / 71 Acheulean hand axe (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 Mousterian tools I Associated with Homo ergaster/erectus I 1.5–0.3 myr ago I 300–30 kya I Essentially unchanged for 1.2 myr I Europe and I Wide distribution in space western Asia 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 Solutrean tools I 19–15 kya Aurignacian tools I Laurel-leaf I 34–23 kya points I Earliest Upper I Later Upper Paleolithic, Europe Paleolithic, I Modern humans Europe I Modern humans 13 / 71 14 / 71 LASCAUX 17 kya Grotte Chauvet 31 kya 15 / 71 16 / 71 Art Venus of Deer’s head Lespugue 18–10 kya (28 kya) 17 / 71 18 / 71 Spear 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 Tool 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 Aurignacian I Each tradition occupies a I gracile 28–22 kya Gravettian small region. 19–15 kya Solutrean I flat face I Culture varies in space. 18–10 kya Magdalenian I chin I forehead I domed skull 21 / 71 22 / 71 Herto, Ethiopia (160 kya) Manot, Israel (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, Basques: late expansions Africa, Europe, East Asia, and India. I Eurasian populations left Africa about 100 thousand years ago Paleolithic or Neolithic? 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 Siberia 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 Pleistocene bottleneck. 35 / 71 36 / 71 Neanderthal 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, Romania 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 Younger Dryas). (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 Neolithic Europe 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 Ancient North Eurasian 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, light skin, 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 art, blade 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 71 / 71.