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Home , Archaic humans

The Fate of the Announcement

The first midterm exam will take place in class on Monday, October 1. You only need to bring a pencil or pen with you.

The exam will have 2 parts: 1) 16 multiple choice questions (1 point each, 15 minutes) 2) 6 out of 7 short IDs (4 points each, 30 minutes)

Sample questions are posted on the course website. After erectus, several species of archaic Homo emerged during the

Homo heidelbergensis, Neanderthals, and are often called “Archaic Homo” or “Archaic ”

Neanderthals Homo sapiens Denisovans (400-30kya) (200kya-present) (40kya) Europe and Middle East first Asia

Homo heidelbergensis (600-200kya) Africa, Europe, and the Middle East

Homo erectus (2mya-150kya?) Africa and Asia. Europe? Outline of Today’s Class: The Fate of the Neanderthals

1) biology and behavior

2) Modern humans - Early expansions out of Africa >100kya - Later expansions out of Africa 60kya associated with modern behavior

3) Neanderthal ancient DNA and clues about The Neanderthals (300kya-30kya) • First fossils discovered in 1856 in the Neander Valley, Germany. • Large brains, large noses, and large brow ridges. Short, stocky bodies. • Cold-weather adapted to life during the “Ice Ages” (ca. 736kya-11.7kya). • Covered a wide geographic range extending from Europe to the Near Boyd & Silk (2014). East and West Asia. • Neanderthals living in different regions had different diets and behaviors (e.g., European Neanderthals hunted large game whereas Near Eastern Neanderthals had a more diverse diet of plants and

small animals). http://www.sciencemag.org/site/special/neandertal/feature/index.html Neanderthal Pathologies and Behavior

• Skeletons often have pathologies such as broken bones or missing teeth. • Patterns of bone breakage are most similar to modern rodeo riders (Berger and Trinkhaus 1995). Neanderthals probably engaged in close quarters hunting. • Many Neanderthal individuals had serious disabilities. Evidence for care for the elderly and disabled. • Possible evidence for intentional burial of the dead at Shanidar , Israel.

Neanderthal skull with missing teeth, La Chapelle-aux-Saints, France Artist reconstruction of Shanidar cave burial, Israel

Berger, T. D., & Trinkaus, E. (1995). Patterns of trauma among the Neandertals. Journal of Archaeological Science, 22(6), 841-852. Industries When did humans first migrate out of Africa?

• Growing evidence for an early migration 100kya via a “southern route”: – Sites in the Arabian peninsula and South Asia contain stone tools dating to 125kya. These tools have stylistic similarities to those made my modern humans in North Africa at the same date. – Skhul and Qafzeh in Israel contain Homo sapiens skeletons dated to 100kya. These are the earliest Homo sapiens remains outside of Africa. Early human skull from Skhul, Israel. – Human teeth and bones have been found at Daoxian (80-120kya) and Zhirendong (>100kya) in southern China. – Genetic evidence that there may have already been an early Homo sapiens population present in Southeast Asia when later migrants arrived.

• Early migrations out of Africa may not have been successful. No evidence for fully “modern” human behavior during these first migrations.

Liu, W., et al. (2015). The earliest unequivocally modern humans in southern China. 526: 696-699. A Second Human Migration Out of Africa 70-50kya The major population expansion out of Africa is associated with fully “modern” human behavior. Modern Human Behavior Emerged Gradually in Africa

Early evidence for modern human behavior at , South Africa (100-75kya): – Shell beads – Ochre for painting bodies – Rudimentary carvings and decorations

Shell Beads, Blombos (77kya)

Processing ochre in abalone shells, Blombos (100kya) Carved ochre, Blombos (75kya)

d'Errico, F., et al. (2005). Nassarius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle . Journal of , 48(1), 3-24. Henshilwood, C. S., et al. (2009). Engraved ochres from the levels at Blombos Cave, South Africa. Journal of human evolution, 57(1), 27-47. Henshilwood, C. S., et al. (2011). A 100,000--old ochre-processing workshop at Blombos Cave, South Africa. science, 334(6053), 219-222. Stone Tool Industries Upper Paleolithic Tools Associated with Homo sapiens

Antler Tools

Upper Paleolithic Stone Tools Indirect Percussion Technique Bone Needles for Making Blades

Atlatl/ Thrower Upper Paleolithic Symbolic Behavior and Art

Painted stones, Apollo 11, Namibia

Mammoth ivory flute, Cave paintings, Chauvet, France Geißenklösterle, Germany

“Venus of Willendorf” figurine, Willendorf, Austria

Cave paintings, , France “Lion Man” figurine carved from Carved atlatl mammoth ivory, Stadel, Germany La Madeleine , France What caused the extinction of the Neanderthals?

Neanderthals were less able to adapt to changing environments. Modern humans may have pushed them into more remote regions until they ultimately went extinct 30,000 ago. Ancient DNA shows that Neanderthals and modern humans interbred

• Initial mtDNA studies indicated that there was no admixture with humans.

• mtDNA is not sufficient for making claims about admixture. Nuclear DNA is always needed.

• Nuclear DNA studies now show that there was admixture between Neanderthals and humans (Green et al. 2010; Prufer et al. 2014). – All non-Africans derive ~2% ancestry from Neanderthals. – Admixture took place in the Near East or South Asia before East Asian and West Asian modern human populations split.

Admixture occurred when humans spread into the Neanderthal’s range in the Near East ca. 54-49kya. As humans continued to expand into other parts of Eurasia, they brought Neanderthal ancestry with them.

Green, R. E., et al. (2010). A draft sequence of the Neandertal . science, 328(5979), 710-722. http://www.sciencemag.org/site/special/neandertal/feature/index.html Prüfer, K. et al. (2014). The complete genome sequence of a Neandertal from the Altai Mountains. Nature, 505(7481), 43. Other Discoveries from Neanderthal Ancient DNA

• Neanderthals have FOXP2 gene allele associated with language (Krause et al. 2007).

• Neanderthals probably had light skin and red or blonde (Lalueza-Fox et al. 2007).

• Neanderthals have extremely low genetic variation across populations living in different geographic regions. Neanderthals probably had a very small population size (Prufer et al. 2014).

Krause, J., et al. (2007). The derived FOXP2 variant of modern humans was shared with Neandertals. Current biology, 17(21), 1908-1912. Lalueza-Fox, C., et al. (2007). A melanocortin 1 receptor allele suggests varying pigmentation among Neanderthals. Science, 318(5855), 1453-1455. Prüfer, K. et al. (2014). The complete genome sequence of a Neandertal from the Altai Mountains. Nature, 505(7481), 43. Other Discoveries from Neanderthal Ancient DNA

• A Neanderthal from Denisova Cave, Altai Mountains, Siberia (Prufer et al. 2014):

– Ancient DNA from a finger bone that belonged to a Neanderthal female.

– DNA shows inbreeding between her close relatives. Her parents were either half siblings, double first cousins, an uncle and a niece, an aunt and a nephew, a grandfather and a granddaughter, or a grandmother and a grandson.

– The Altai Neanderthals were a very isolated, small population (about 600 people 50kya).

Prüfer, K. et al. (2014). The complete genome sequence of a Neandertal from the Altai Mountains. Nature, 505(7481), 43. What caused the extinction of the Neanderthals?

Neanderthals were less able to adapt to changing environments. Modern humans may have pushed them into more remote regions until they ultimately went extinct 30,000 years ago. Next time…Denisovans and Hobbits!