Outline. Early Modern Humans. Moderns invade Eurasia. Acheulean hand axe ( mya) Oldowan tools mya

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1 Outline Early Modern Humans Alan R. Rogers February 7, 2018 Archaeology and paleontology Expansion out of Africa Paleolithic Eurasia Mesolithic Eurasia 1 / 71 2 / 71 Moderns invade Eurasia Oldowan tools mya Earliest stone tools Probably made by Homo habilis/rudolphensis 3 / 71 4 / 71 Acheulean hand axe ( mya) Oldowan tools mya 5 / 71 6 / 71

2 Acheulean tools ( mya) Distribution of Acheulean tools 7 / 71 8 / 71 Acheulean tools Mousterian tools Associated with Homo ergaster/erectus myr ago Essentially unchanged for 1.2 myr Wide distribution in space kya Europe and western Asia Neandertals 9 / / 71 Neandertal sites Mousterian tools Neandertals kya Little change in 270 kyr Wide distribution in space 11 / / 71

3 Solutrean tools Aurignacian tools kya Earliest Upper Paleolithic, Europe Modern humans kya Laurel-leaf points Later Upper Paleolithic, Europe Modern humans 13 / / 71 LASCAUX 17 kya Grotte Chauvet 31 kya 15 / 71 Art Deer s head kya 17 / / 71 Venus of Lespugue (28 kya) 18 / 71

4 Spear thrower increases force of throw penetrated Spanish armor kill at 70 m by 14 kya NTAL UPPER PALE- OLTHC OF EUROPE 19 / / 71 Tool traditions of the Upper Paleolithic Age Tradition kya Chatelperronian kya Aurignacian kya Gravettian kya Solutrean kya Magdalenian Change is rapid. Each tradition occupies a small region. Culture varies in space. Neandertal & Modern Modern (at right): small long-limbed gracile flat face chin forehead domed skull 21 / / 71 Herto, Ethiopia (160 kya) Manot, srael (55 kya) Modern But with archaic features, such as occipital bun. 23 / / 71

5 Outline Comas et al (1997) studied European mismatch distributions Archaeology and paleontology Expansion out of Africa Paleolithic Eurasia Mesolithic Eurasia 25 / / 71 Mismatch distributions suggest expansion across Europe Nuclear DNA: Xing et al 2010 Mid-East and Turkey: early expansions. British, Basques: late expansions Paleolithic or Neolithic? Studied large sample of nuclear DNA from populations in Africa, Europe, East Asia, and ndia. Eurasian populations left Africa about 100 thousand years ago Did not separate until thousand years ago Comas et al thought Paleolithic but may have been misled by mtdna clock. 27 / / 71 Outline Archaeology and paleontology Expansion out of Africa Paleolithic Eurasia Mesolithic Eurasia 29 / / 71

6 Paleolithic DNA of NW Eurasia Ust -shim: a 45-ky-old modern man from W Siberia Excavated 2008 Fu et al. (2014) Note time scale at back. 31 / 71 Location of Ust -shim 32 / 71 Dates and temperature 1, Ust -shim; 4, Denisova. 33 / / 71 History of population size Ust -shim PC map Ust -shim (purple) similar to Central Asians Red, assumes Ust -shim lived today; Blue, fits curve to others and estimates mutation rate. Lived during late Pleistocene bottleneck. 35 / / 71

7 Neanderthal allele sharing Conventional wisdom Early emigration out of Africa to Australia & New Guinea. Later northern emigration. Ust -shim should be part of northern population. Problem Ust -shim is equally related to northern and southern modern populations. Perhaps it was a 3-way split, and Ust -shim s population went extinct. Ust -shim has 2.3% Neanderthal DNA in big chunks. mplies recent admixture (50 60 kya). 37 / / 71 Man from Peştera cu Oase, Romania Oase 1 40 ky-old 6 9% of genome is Neanderthal 3 larger than any other 3 Neanderthal segments >50 cm mplies 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 Kostenki 14: a 37 ky old Russian man Affinities of Kostenki with world populations Similar to Northern Europeans. (Seguin-Orlando et al. 2014) 41 / / 71

8 Affinities of Kostenki with European populations Neandertal allele sharing (Seguin-Orlando et al. 2014) Similar to northern and western Europeans. (Seguin-Orlando et al. 2014) K14 has more Neandertal DNA and in bigger chunks. 43 / / 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. mplies recent admixture 54 kya. (Seguin-Orlando et al. 2014) (Raghavan et al. 2013) 45 / / 71 Location of Mal ta site Mal ta in principal components map Raghavan et al. (2013) 47 / 71 (Raghavan et al. 2013) 48 / 71

9 Affinities of Mal ta with other populations 24 ky old burial from Mal ta, Siberia 1/3 of ancestry shared with Native Americans and Europeans European mitochondrial DNA (Raghavan et al. 2013) (Raghavan et al. 2013) Similar to Amerindians and Northern Europeans. 49 / 71 mtdna haplogroups: kya 50 / 71 mtdna haplogroups: kya (Posth et al 2016) (Posth et al 2016) 51 / 71 mtdna haplogroups: kya 52 / 71 mtdna haplogroups: kya (Posth et al 2016) (Posth et al 2016) 53 / / 71

10 Outline Two big changes in variation. One coincides with last glacial maximum (LGM). The other with the last glacial (aka the Younger Dryas). Archaeology and paleontology Expansion out of Africa Paleolithic Eurasia Mesolithic Eurasia (Posth et al 2016) 55 / / 71 La Braña: a 7,000 y old forager La Braña site 57 / / 71 La Braña PC Map: World La Braña PC Map: Europe On world-wide scale, La Braña is European. But La Braña is not like modern Europeans. 59 / / 71

11 Neandertal admixture fraction Gradual decline in Neanderthal admixture La Braña had more Neanderthal DNA than modern humans. (Seguin-Orlando et al. 2014) 61 / / 71 La Braña adaptations Study of Lazaridis et al (2014) Dark skin Blue eyes Lactose intolerant Poor at digesting starch. Modern allele at 24 of 40 loci affecting immune function. These observations suggest that the Neolithic transition did not drive all cases of adaptive innovation on immunity genes found in modern Europeans. (Olalde et al 2014) DNA sequences of: Label Location Samples Age Description Loschbour Luxembourg 1 8 ky forager Motala Sweden 7 8 ky forager Stuttgart Germany 1 7 ky farmer Europe ky modern The moderns were not sequenced but were genotyped at 594,924 autosomal single-nucleotide polymorphisms. 63 / / 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

12 Contributions to modern Europeans PCA map ANE WHG EEF Ancient North Eurasian West-European hunter-gatherer Early European Farmer 67 / / 71 Contributions to modern Europeans Adaptations Lochbour (forager) relatively good at digesting starch, even before agriculture. Stuttgart (farmer): Dark hair, light skin, brown eyes. Foragers: Dark skin, hair and blue eyes. But one Motala (forager) sample had one copy of light skin allele. t was in Europe before agriculture. 69 / / 71 Summary Archaics changed slowly both in time and space. Pace of change is faster for modern humans. Moderns appear first in Africa, 160 ky ago. Spread through Eurasia 50 ky ago. ntroduce art, blade tools, projectile weapons. Genome of Ust -shim, 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. Kostenki, a 36 ky-old Siberian, genetically similar to Europeans. More Neanderthal DNA than modern humans. Mal ta 71 / 71