Tohoku Forum for Creativity
Thematic Program 2022

Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology,
Palaeoanthropology, and Genetics
September 27 – October 4, 2022
Sendai, JapanPROGRAM AND ABSTRACTS

Edited by
Katsuhiro Sano & Waka KuboyamaTohoku Forum for Creativity Thematic Program 2022 “Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology, Palaeoanthropology, and Genetics”
Editor-in-Chief
Katsuhiro Sano, Tohoku University
Editorial Board
Katsuhiro Sano, Tohoku University
Waka Kuboyama, Tohoku University
Masami Izuho, Tokyo Metropolitan University
Kohei Tamura, Tohoku University
Stefano Benazzi, University of Bologna
Published by
Tohoku Forum for Creativity
Tohoku University
2-1-1 Katahira, Aoba-ku, Sendai, 980-8577, JAPANTEL: +81-22-217-6292
creativity@grp.tohoku.ac.jp
https://www.tfc.tohoku.ac.jp
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in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the priorpermission of the Tohoku Forum for Creativity.
Cover image: Miho Iwabuchi© Tohoku Forum for Creativity

CONTENTS
Preface……………………………….…………………………….……..………..………….…..…..i
Program ...………………………...………………………………….…..…………………..….……ii
International Symposium
Insights Into Human History in the Eurasian Stone Age: Recent Developments in Archaeology,
Palaeoanthropology, and Genetics
Session 1: Migration of archaic humans into Central and Northeast Asia………………...……….…1
Kseniya A. Kolobova
Neanderthal migration to the East ………………………….………………………………………...2
Viviane Slon
Genetic studies on Denisovans and Neanderthals ……………..………………………………….....3
Viviane Slon
Inferences on the genetic history of Eurasia based on sedimentary ancient DNA…………………..4
Olaf Jöris & Marcel Weiss
The spatio-temporal patterning of Keilmessergruppen assemblages: Implications for Neanderthal
dispersal and migrations ……………………………………………………….………….….……..5
Yoshihiro Nishiaki
Teshik-Tash Neanderthal lithic industry in the context of Neanderthal dispersals……………..……7
Xing Song
Discussion on the evolutionary status of mainland East Asian late Middle Pleistocene archaic
Homo…………………………………….…………………………………………………….…….8
Dongju Zhang et al.
Denisovans from Baishiya Karst Cave on the Tibetan Plateau…………….…………………….….9
Xing Gao
The complexity of the Middle Paleolithic industries in China and implications for trajectories
of human evolution in the region. ………….……………………………………………..………10
Katerina Douka
What we know about Denisovans: An overview of current fossils, archaeology, chronology andgeographic spread………………………….……………………………………….…….…………11
Frido Welker
Palaeoproteomics for human evolution……….……………..……………………………..…….…13

Session 2: Origin and expansion of Homo sapiens in western Eurasia..………….………….…...…15
Jean-Jacques Hublin
Origin and expansion of Homo sapiens in western Eurasia ……………………………….….……16
Israel Hershkovitz
Pre-Neanderthals and modern humans in the Levant during the late Middle Pleistocene:
A perspective from the Nesher Ramla Homo fossil…………………………………………………17
Seiji Kadowaki
Multiple pathways to the Upper Paleolithic? Geographic variability in the Middle to Upper
Paleolithic transition in the Levant………………………………………………………….……...18
Andrea Columbu
MIS3 palaeoclimate and palaeoenvironment in western Eurasia…………………………….…….19
Cosimo Posth
The genomic profile of Upper Paleolithic European hunter-gatherers …………………….………20
Tsenka Tsanova
The Initial Upper Palaeolithic at Bacho Kiro Cave and in the Eastern Balkan area…………….….21
Andrea Picin
Cultural perspectives on early dispersals of Homo sapiens in Central Europe and Eastern
European Plains ……………………………………………………………………………………23
Stefano Benazzi
ERC project SUCCESS: The earliest migration of Homo sapiens in southern Europe……………24
Katsuhiro Sano et al.
Advanced projectile technology of the earliest Homo sapiens in Europe …………………………25
Sahra Talamo
What a precise clock! ERC project RESOLUTION provides more refine chronology of theInitial and Early Upper Palaeolithic in Europe……………..………………………………..……..26
Special Session: The Japanese Palaeolithic: Isolation and cultural exchanges ………………………29
Yousuke Kaifu
Another Palaeolithic frontier: Modeling the earliest seafaring in East Asia………………………..30
Katsuhiro Sano et al.
The spatio-temporal patterns of early Upper Palaeolithic assemblages in the Japanese islands …..31
Kazuki Morisaki
Middle and late Upper Palaeolithic in the Japanese archipelago: Local development and continental
influence ……………..……………………………………………………………………………..32
Fumie Iizuka
Ecosystem change and decisions for pottery using foraging: Late Pleistocene cases from the Japanesearchipelago ……………………………………………………………….…………….…………..33
Hiroki Oota
Ancient genome analysis of human remains in the Japanese archipelago…………………………34
Session 3: The earliest migration of Homo sapiens into Central and Northeast Asia………………..37
Ted Goebel
Problems and prospects in the study of modern-human dispersal in Northern Asia…………….…38
Masami Izuho
Early adaptations into tundra landscape by modern humans: A case study of the Initial
Upper Paleolithic in Mongolia and Transbaikal, Russia ………………………….…………….….39
Nicolas Zwyns
Exploring the variability of IUP assemblages using a techno-economic approach: The example ofTolbor-16, Mongolia ……………………………………………………………………….……....40
Evgeny Rybin et al.
Initial Upper Paleolithic of Southern Siberia and Central Asia: Continuity and variability …….…41
Feng Li
Understanding the Initial Upper Paleolithic of China in a regional context of Eastern Eurasia…...42
Fei Peng et al.
New progress on the excavation and research of Shuidonggou site complex……………………...43
Xiaowei Mao
Genetic insight into northern East Asia during the period of Ice Age and Paleolithic-Neolithic
transition………………………………………………………….………………………………...44
Steven Kuhn
The Initial Upper Paleolithic in Asia: Is it still a useful concept?.....................................................45
Tom Higham
Pitfalls and progress in dating the earliest appearance of Homo sapiens in Central and Northeast
Asia………………………………………………………………………………..………...…...…46
Workshop 1: Emergence of regional diversity of Northeast Asia…………………..………….…….47
Kelly Graf
Regional differentiation from EUP to LUP in Northeast Asia: An overview………..……………..48
Arina Khatsenovich et al.
Upper Paleolithic exchange networks in Siberia and Mongolia……………………..………….….49
Shejiang Wang
Paleolithic archaeology in the Qinling Mountains region, Central China…………..………….…..50
Olaf Jöris

The ebb and flow of human dispersals into Central Europe and further North from the Last Glacial
Maximum to the beginning of Late Glacial warming, 25–14 cal kBP…………………….……...….51
Maxim Kozlikin
Denisovans: Age, culture and habitat.………….…………………………………………….……..53
Anton Anoikin et al.
10,000 years without Levallois: IUP industries of Ushbulak site, Eastern Kazakhstan……………54
Workshop 2: Recovering ancient remains and reconstructing past………………………….………55
Simona Arrighi
Function and behaviour: Recognizing use-wear on prehistoric tools …………………………….56
Yoshitaka Kanomata
Methodology for the use-wear analysis developed by Tohoku University Microwear Research
Team (TUMRT).……...…………………………………………………………………………….57
Shunsuke Totsuka & Katsuhiro Sano
Methods for identifying hunting traces and its application for early Upper Palaeolithic sites in Japan…….……………………………………………………………………………………………..…58
Kohei Tamura
Quantifying morphological variation: Applications of geometric morphometrics to archaeologicalartifacts……………………………………………………………………………………….……..59
Matteo Rossini & Clarissa Dominici
Approaching Upper Palaeolithic hunting strategies through technology and residue analysis….…60
Francesco Boschin & Jacopo Crezzini
Zooarchaeology and the Middle to Upper Palaeolithic transition: Some methodological insights…61
Vincenzo Spagnolo
Disentangling the spatial structure of a Palaeolithic camp: Methodological issues………………..62
Giulia Marciani
Lithic technology, the notion of chaîne opératoire and its application in two case studies: The
Mousterian of Riparo l’Oscurusciuto and the Uluzzian of Grotta di Castelcivita……………….…63
Federico Lugli
Elemental and isotopic analyses of tooth enamel disclose individuals’ life histories………………..64
Author Index…………………...…………….………………………………………………………65
Memo……………………………………………………………….……………………….……….67

PREFACE
We are delighted to welcome you to this historical city of Sendai, Japan, for the Tohoku Forum for
Creativity (TFC) Thematic Program 2022 titled “Insights Into Human History in the Eurasian Stone
Age: Recent Developments in Archaeology, Palaeoanthropology, and Genetics.” The TFC at Tohoku
University is the first international visitor research institute in Japan. The forum aims to identify issues
across all fields of research, and develop innovative ideas and deeper theoretical foundations throughintensive, focused discussions.
It is now well-known that Neanderthals and Denisovans lived in Eurasia, before the expansion of
Homo sapiens. Although Neanderthals and Denisovans were genetically and culturally close to Homosapiens, they became extinct after the arrival of Homo sapiens. In contrast, the population size of
Homo sapiens steadily increased. However, there is limited information on how Homo sapiens
increased their population size, while the other human species became extinct. An important goal ofthis thematic program is to share recent advances in archaeology, palaeoanthropology, and genetics.
Our TFC Thematic Program 2022 comprises an international symposium and two international
workshops, and incorporates 50 oral presentations, including five keynote lectures and a special
lecture. It is a great honor for us to host comprehensive presentations on the Eurasian Stone Age andwelcome world-class researchers who have contributed to the field through cutting-edge studies. We
sincerely appreciate all speakers for their presentations, which enable us to hold this thematic program.
In particular, we would like to express our deepest gratitude to our colleagues who came from overseas
despite difficult situations. We would also like to thank all participants who arrived in person as wellas online.
The international symposium/workshops is supported by TFC and the Center for Northeast AsianStudies (CNEAS), Tohoku University. We thank the staff of TFC and CNEAS for their support. Lastly,
we are particularly grateful to the secretariat Waka Kuboyama (CNEAS) and program coordinator
Takao Koga (TFC), without whose support this event would not have been possible.
With best wishes,
Principal organizer

Katsuhiro Sano
CNEAS, Tohoku UniversityiINTERNATIONAL SYMPOSIUM
Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology, Palaeoanthropology, and Genetics
Date: September 27 – 29, 2022
Venue: TOKYO ELECTRON House of Creativity, Tohoku University
Organizers: Katsuhiro Sano, Masami Izuho, Kohei Tamura, & Stefano Benazzi

PROGRAMSEPTEMBER 27 (TUESDAY)
11:00 – 13:00 Registration
13:00 – 13:10 Opening address
Katsuhiro Sano (Deputy Director, Center of Northeast Asian Studies, Tohoku
University)
13:10 – 13:20 Greetings
Akira Ono (Honorary President of Asian Palaeolithic Association/ Emeritus
Professor, Tokyo Metropolitan University)

Session 1: Migration of archaic humans into Central and Northeast Asia
Chairperson: Stefano Benazzi
13:20 – 14:20 Keynote lecture: Neanderthal migration to the East [online]
Kseniya A. Kolobova
14:20 – 14:40 Coffee break
14:40 – 15:00 Viviane Slon
Genetic studies on Denisovans and Neanderthals
15:00 – 15:20 Viviane Slon
Inferences on the genetic history of Eurasia based on sedimentary ancient DNA
15:20 – 15:40 Olaf Jöris & Marcel Weiss
The spatio-temporal patterning of Keilmessergruppen assemblages: Implications for
Neanderthal dispersal and migrations

15:40 – 16:00 Yoshihiro Nishiaki
Teshik-Tash Neanderthal lithic industry in the context of Neanderthal dispersals

16:00 – 16:20 Coffee break
16:20 – 16:40 Xing Song
Discussion on the evolutionary status of mainland East Asian late Middle Pleistocene
archaic Homo [online]ii

16:40 – 17:00 Dongju Zhang, Frido Welker, Chuan-Chou Shen, Bo Li, Jean-Jacques
Hublin, Svante Pääbo, Qiaomei Fu, & Fahu Chen
Denisovans from Baishiya Karst Cave on the Tibetan Plateau [online]
17:00 – 17:20 Xing Gao
The complexity of the Middle Paleolithic industries in China and implications for
trajectories of human evolution in the region [online]
17:20 – 17:40 Katerina Douka
What we know about Denisovans: An overview of current fossils, archaeology,
chronology and geographic spread [online]
17:40 – 18:00 Frido Welker
Palaeoproteomics for human evolution [online]
18:00 – 18:10 Coffee break
18:10 – 18:50 Discussion

SEPTEMBER 28 (WEDNESDAY)
08:50 – 9:00

Greetings
Motoko Kotani (Executive Vice President for Research, Director of Tohoku Forumfor Creativity, Tohoku University)

Session 2: Origin and expansion of Homo sapiens in western Eurasia
Chairperson: Masami Izuho
09:00 – 10:00 Keynote lecture: Origin and expansion of Homo sapiens in western Eurasia
Jean-Jacques Hublin
10:00 – 10:20 Coffee break
10:20 – 10:40 Israel Hershkovitz
Pre-Neanderthals and modern humans in the Levant during the late Middle
Pleistocene: A perspective from the Nesher Ramla Homo fossil [online]
10:40 – 11:00 Seiji Kadowaki
Multiple pathways to the Upper Paleolithic? Geographic variability in the Middle toUpper Paleolithic transition in the Levant
11:00 – 11:20 Andrea Columbu
MIS3 palaeoclimate and palaeoenvironment in western Eurasia
11:20 – 11:40 Cosimo Posth
The genomic profile of Upper Paleolithic European hunter-gatherers
11:40 – 12:00 Tsenka Tsanova
The Initial Upper Palaeolithic at Bacho Kiro Cave and in the Eastern Balkan area
[online]iii

12:00 – 13:30 Lunch
13:30 – 13:50 Andrea Picin
Cultural perspectives on early dispersals of Homo sapiens in Central Europe andEastern European Plains
13:50 – 14:10 Stefano Benazzi
ERC project SUCCESS: The earliest migration of Homo sapiens in southern
Europe14:10 – 14:30 Katsuhiro Sano, Simona Arrighi, Lisa Vaccari, Stefano Benazzi, & Adriana Moroni
Advanced projectile technology of the earliest Homo sapiens in Europe
14:30 – 14:50 Sahra Talamo
What a precise clock! ERC project RESOLUTION provides more refine chronologyof the Initial and Early Upper Palaeolithic in Europe [online]
14:50 – 15:00 Coffee break
15:00 – 15:40 Discussion

Special Session: The Japanese Palaeolithic: Isolation and cultural exchanges
Chairperson: Kohei Tamura
16:00 – 16:40 Special lecture: Another Palaeolithic frontier: Modeling the earliest seafaring
in East Asia
Yousuke Kaifu
16:40 – 17:00 Katsuhiro Sano, Shunsuke Totsuka, Masami Izuho, & Kazuki Morisaki
The spatio-temporal patterns of early Upper Palaeolithic assemblages in the Japaneseislands
17:00 – 17:20 Kazuki Morisaki
Middle and late Upper Palaeolithic in the Japanese archipelago: Local development
and continental influence
17:20 – 17:40 Fumie Iizuka
Ecosystem change and decisions for pottery using foraging: Late Pleistocene cases
from the Japanese archipelago
17:40 – 18:00 Hiroki Oota
Ancient genome analysis of human remains in the Japanese archipelago
18:00 – 18:10 Coffee break
18:10 – 18:40 Discussion

iv

SEPTEMBER 29 (THURSDAY)
Session 3: The earliest migration of Homo sapiens into Central and Northeast Asia
Chairperson: Katsuhiro Sano
09:30 – 9:50

Ted Goebel
Problems and prospects in the study of modern-human dispersal in Northern Asia

9:50 – 10:10

Masami Izuho
Early adaptations into tundra landscape by modern humans: A case study of the Initial
Upper Paleolithic in Mongolia and Transbaikal, Russia

10:10 – 10:30 Nicolas Zwyns
Exploring the variability of IUP assemblages using a techno-economic approach: The
example of Tolbor-16, Mongolia
10:30 – 10:50 Coffee break
10:50 – 11:10 Evgeny Rybin, Kseniya Kolobova, & Arina Khatsenovich
Initial Upper Paleolithic of Southern Siberia and Central Asia: Continuity andvariability
11:10 – 11:30 Feng Li
Understanding the Initial Upper Paleolithic of China in a regional context ofEastern Eurasia [online]
11:30 – 11:50 Fei Peng, Huimin Wang & Xing Gao
New progress on the excavation and research of Shuidonggou site complex [online]
11:50 – 12:10 Xiaowei Mao
Genetic insight into northern East Asia during the period of Ice Age and PaleolithicNeolithic transition [online]
12:10 – 14:00 Lunch
14:00 – 14:20 Steven Kuhn
The Initial Upper Paleolithic in Asia: Is it still a useful concept?
14:20 – 15:20

Keynote lecture: Pitfalls and progress in dating the earliest appearance of
Homo sapiens in Central and Northeast Asia [online]
Tom Higham

15:20 – 16:00 Discussion

Poster Session 16:30 – 18:00

v

WORKSHOP 1
Emergence of regional diversity of Northeast Asia
Date: September 30, 2022
Venue: TOKYO ELECTRON House of Creativity, Tohoku University
Organizers: Masami Izuho & Katsuhiro Sano

PROGRAM
09:00 – 9:40 1st keynote lecture: Regional differentiation from EUP to LUP in Northeast Asia:
An overview
Kelly Graf
09:40 – 10:00 Arina Khatsenovich, Rybin E., Shelepaev R.A., & Olsen J.W.
Upper Paleolithic exchange networks in Siberia and Mongolia [online]
10:00 – 10:20 Shejiang Wang
Paleolithic archaeology in the Qinling Mountains region, Central China [online]
10:20 – 10:40 Coffee break
10:40 – 11:20 2nd keynote lecture: The ebb and flow of human dispersals into Central
Europe and further North from the Last Glacial Maximum to the beginning of
Late Glacial warming, 25–14 cal kBP
Olaf Jöris
11:20 – 11:40 Maxim Kozlikin
Denisovans: Age, culture and habitat [online]
11:40 – 12:00 Anton Anoikin, Kharevich Vladimir, Pavlenok Galina, & Taimagambetov Zhaken
10,000 years without Levallois: IUP industries of Ushbulak site, Eastern Kazakhstan[online]12:00 – 12:20 Coffee break
12:20 – 13:00 Discussion
Museum Tour of SENDAI CITY TOMIZAWA SITE MUSEUM 15:00 – 16:45

vi

WORKSHOP 2
Recovering ancient remains and reconstructing past
Date: October 4, 2022
Venue: TOKYO ELECTRON House of Creativity, Tohoku University
Organizers: Stefano Benazzi, Adriana Moroni, & Katsuhiro Sano

PROGRAM
13:30 – 13:50 Opening address and introduction
Stefano Benazzi
13:50 – 14:10 Simona Arrighi
Function and behaviour: Recognizing use-wear on prehistoric tools
14:10 – 14:30 Yoshitaka Kanomata
Methodology for the use-wear analysis developed by Tohoku University Microwear
Research Team (TUMRT)
14:30 – 14:50 Shunsuke Totsuka & Katsuhiro Sano
Methods for identifying hunting traces and its application for early Upper
Palaeolithic sites in Japan
14:50 – 15:10 Kohei Tamura
Quantifying morphological variation: Applications of geometric morphometrics toarchaeological artifacts
15:10 – 15:30 Coffee break
15:30 – 15:50 Matteo Rossini & Clarissa Dominici
Approaching Upper Palaeolithic hunting strategies through technology and residue
analysis [online]
15:50 – 16:10 Francesco Boschin & Jacopo Crezzini
Zooarchaeology and the Middle to Upper Palaeolithic transition: Somemethodological insights [online]
16:10 – 16:30 Vincenzo Spagnolo
Disentangling the spatial structure of a Palaeolithic camp: Methodological issues[online]
16:30 – 16:50 Giulia Marciani
Lithic technology, the notion of chaîne opératoire and its application in two casestudies: The Mousterian of Riparo l’Oscurusciuto and the Uluzzian of Grotta diCastelcivita [online]
16:50 – 17:10 Federico Lugli
Elemental and isotopic analyses of tooth enamel disclose individuals’ life histories[online]17:10 – 17:30 Coffee break
17:30 – 18:30 Discussionvii

INTERNATIONAL SYMPOSIUM
Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology, Palaeoanthropology, and Genetics

Session 1
Migration of archaic humans into
Central and Northeast Asia

INTERNATIONAL SYMPOSIUM

Keynote Lecture

Neanderthal migration to the East________________________________________________________________________________

Kseniya A. Kolobova
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences________________________________________________________________________________
Neanderthals were once widespread across Europe and western Asia. They also penetrated into the
Altai Mountains of southern Siberia, but the geographical origin of these populations and the timingof their dispersal have remained elusive. Here we present archaeological and anthropologicalassemblages from Chagyrskaya, Okladnikov and Strashnaya Caves at Altai represented late
Neanderthals population between 59 and 44 thousand years ago. Environmental reconstructions
suggest that the hominins were adapted to the dry steppe and hunted bison/horses. Their distinctive
toolkit closely resembles Micoquian/Keilmessergruppen (KMG) assemblages from central and eastern
Europe, including the northern Caucasus, more than 3,000 kilometers to the west of Altai. We identify
eastern Europe as the most probable ancestral source region for the Altai toolmakers, supported by
DNA results linking the Neanderthal remains with populations in northern Croatia and the northern
Caucasus, and providing a rare example of a long-distance, intercontinental population movement
associated with a distinctive Paleolithic toolkit. Comparative studies of Micoquian archaeologicalcomplexes from Central, Eastern Europe, the Crimea, the Volga region and the Altai havedemonstrated a single pattern of bifacial manufacture. At the moment all Micoquian complexes show
a certain variability, which does not depend on the geographical localization of the sites. Studies ofadaptation behavior suggest that Neanderthals applied strategies known to them in newly occupied
territories. The only recorded difference has to do with the adaptation of lithic technology to the shapeof stone raw materials. The question of the possible migration of Neanderthals to more eastern
territories remains unresolved. There have already been several hypotheses about the discovery of
Neanderthal complexes in Tuva and Kazakhstan. We are also examining the question of a possible
migration to China.
Acknowledgments
The study had been supported by Science Foundation, Project No. 21-18-00376.

2

INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Genetic studies on Denisovans and Neanderthals________________________________________________________________________________

Viviane Slon1,2
1) Department of Anatomy and Anthropology and Department of Human Molecular Genetics and Biochemistry2) The Dan David Center for Human Evolution and Biohistory Research, Sackler Faculty of Medicine, Tel Aviv
University________________________________________________________________________________
Genetic research conducted on ancient hominins can enrich our understanding of our own evolutionary
history and that of our closest extinct relatives, the Neanderthals and the Denisovans, yielding insights
on demography, population origins and admixture events. While Neandertals, who lived in western
Eurasia, are well-characterized morphologically based on numerous remains in the fossil record, only
a handful of remains have been identified as pertaining to the Denisovans, who are thought to have
inhabited large parts of eastern Eurasia. Here I will summarize recent ancient DNA-based studies
aimed at gaining further knowledge on both groups of archaic hominins, conducted while developing
techniques to circumvent the scarcity of ancient skeletal remains available for genetic analyses.

3

INTERNATIONAL SYMPOSIUM

Inferences on the genetic history of Eurasia based on sedimentary
ancient DNA________________________________________________________________________________

Viviane Slon1,2
1) Department of Anatomy and Anthropology and Department of Human Molecular Genetics and Biochemistry2) The Dan David Center for Human Evolution and Biohistory Research, Sackler Faculty of Medicine, Tel Aviv
University________________________________________________________________________________In the past years, DNA retrieved from ancient human remains have been instrumental in furthering
our understanding of our own evolutionary past, as well as that of our closest relatives, the Neandertals
and the Denisovans. Such studies, however, are inherently limited to sites and timeframes where suchremains have been found and made available for sampling. A complementary approach can be torecover human DNA fragments from ancient sediments – a source material ubiquitously andabundantly found at any archaeological excavation. Here, I will outline a methodology to do so,
including its highlights and pitfalls, and demonstrate the potential of this research to shed light on thegenetic history of ancient populations using examples from prehistoric sites in Europe and Asia.

4

INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

The spatio-temporal patterning of Keilmessergruppen Assemblages:
Implications for Neanderthal dispersal and migrations________________________________________________________________________________

Olaf Jöris1,2,3 and Marcel Weiss4
1)

Römisch-Germanisches Zentralmuseum – MONREPOS Archaeological Research Centre and Museum for
Human Behavioural Evolution

2)

Institute of Ancient Studies, Department of Prehistoric and Protohistoric Archaeology, Johannes Gutenberg
University

3)

MOE Key Laboratory of Western China’s Environmental Systems Research School of Arid Environment andClimate Change

4)

Institut für Ur-und Frühgeschichte, Friedrich-Alexander-Universität Erlangen-Nürnberg________________________________________________________________________________The Late Middle Palaeolithic of Central Europe is characterized by an amalgam of different
technological and typological features, hampering a straightforward interpretation of lithic
assemblage variability (Weiss et al. 2017). However, certain techno-typological characteristics
show Regionalised Cultural Signatures (RCS) and allow for the definition of spatio-temporally
defined techno-complexes (Jöris et al. 2022). The Keilmessergruppen (KMG) techno-complex(Jöris 2004) features bifacial backed knives – asymmetric cutting tools with a plano-convex crosssection that display a singular acute edge opposite a natural or roughly worked back. These
Keilmesser represent one of the most studied Central European RCS. Detailed technologicalanalyses have highlighted their inherent potential for (multiple) resharpening, showing they were
designed for long use-life cycles (Jöris 2001; Weiss 2020), and that Keilmesser design andmanufacture relied on the generational transmission of the tool concept through different stagesof social learning (Jöris and Uomini 2019). These aspects make the Keilmesser an excellent fossile
directeur for Late Middle Palaeolithic traditions.
Plotted against time, the geographical patterning of KMG assemblages hint at supra-regionalchanges in the dispersal areas of KMG populations. Earlier studies argued for North-to-Southmigrations in Central Europe resulting from adaptations to the flickering of interstadial and stadialconditions (Jöris 2004). New age estimates for KMG sites (Hein et al. 2020; Weiss et al. 2018,
submitted) call for a revision of the KMG record. The present paper introduces a revised modelof KMG dispersal and expansion throughout northern Eurasia relative to palaeoenvironmental
changes. The results of this study formulate predictions on how the observed changes in KMG
dispersal may be archived in Neanderthal aDNA.
References
Hein, M., Weiss, M., Otcherednoy, A., Lauer, T. 2020. Luminescence chronology of the key-Middle Paleolithic site Khotylevo I(Western Russia) - Implications for the timing of occupation, site formation and landscape evolution. Quaternary Science5

INTERNATIONAL SYMPOSIUM

Advances, 2, e100008.
Jöris, O. 2001. Der Spätmittelpaläolithische Fundplatz Buhlen (Grabungen 1966-69). Stratigraphie, Steinartefakte und Fauna desoberen Fundplatzes. Universitätsforschungen zur Prähistorischen Archäologie 73, Habelt, BonnJöris, O. 2004. Zur chronostratigraphischen Stellung der spätmittelpaläolithischen Keilmessergruppen: Der Versuch einer
kulturgeographischen Abgrenzung einer mittelpaläolithischen Formengruppe in ihrem europäischen Kontext. Berichte derRömisch-Germanischen Kommission, 84, 49–153.
Jöris, O., Neruda, P., Wiśniewski, A., Weiss, M. 2022. (in prep.) The Late and Final Middle Palaeolithic of Central Europe and its
contribution to the formation of the Upper Palaeolithic – review and synthesis.
Jöris, O., Uomini, N. 2019. Evidence for Neanderthal Hand Preferences from the Late Middle Palaeolithic Site of Buhlen, Germany:
Insights into Neanderthal Learning Behaviour. In: Nishiaki, Y., Jöris, O. (eds.) Learning Among Neanderthals and Palaeolithic
Modern Humans. Archaeological Evidence. Replacement of Neanderthals by Modern Humans Series. Springer Nature
Singapore, 77–93.
Weiss, M. 2020. The Lichtenberg Keilmesser - it’s all about the angle. PLoS ONE, 15 (10), e0239718.
Weiss, M., Otcherednoy, A., Wiśniewski, A. 2017. Using multivariate techniques to assess the effects of raw material, flaking behavior
and tool manufacture on assemblage variability: An example from the late Middle Paleolithic of the European Plain. Journalof Archaeological Science, 87, 73–94.
Weiss, M., Lauer, T., Wimmer, R., Pop, C.M. 2018. The Variability of the Keilmesser-Concept: a Case Study from Central Germany.
Journal of Paleolithic Archaeology, 1, 202–246.
Weiss, M., Hein, M., Urban, B., Stahlschmidt, M.C., Heinrich, S., Hilbert, Y.H., Power, R.C. von Suchodoletz, H., Terberger, T.,
Böhner, U., Klimscha, F., Veil, St., Breest, K., Schmidt, J., Colarossi, D., Tucci, M., Frechen, M., Tanner, D.C., Lauer, T.
(submitted). Neanderthals in changing environments from MIS 5 to early MIS 4 in northern Central Europe – Integratingarchaeological, (chrono) stratigraphic and paleoenvironmental evidence at the site of Lichtenberg. Quaternary Science Reviews.

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Teshik-Tash Neanderthal lithic industry in the context of Neanderthal
dispersals______________________________________________________________________
Yoshihiro Nishiaki
The University Museum, The University of Tokyo______________________________________________________________________The Teshik Tash cave in Uzbekistan, excavated in 1937 and 1938, is one of the best-known Middle
Palaeolithic sites associated with Neanderthal fossil remains in western Central Asia. Its lithic industry,
although radiometrically sufficiently dated thus far, provides an important insight into the processesof Neanderthal dispersion to this part of Asia, supposedly from the west. Furthermore, a recent analysisof the available lithic materials stored in Uzbekistan conducted by our team revealed a consistent setof techno-typological features, including the production of common blanks from little-prepared cores
rather than Levallois ones, moderate production of elongated blanks, and very common production ofside scrapers, which characterize the Teshik Tash lithic industry. When compared with the Middle
Palaeolithic lithic industries to those in the regions neighboring to the west, they display similarities
with those of the Zagros Mountains. This is not surprising, considering the geographical location of
the Teshik Tash Cave. Rather, their dissimilarities with those located further west, such as in the Levant
and the Caucasus regions, would provide a useful insight to understanding the cultural landscapes of
the Neanderthal communities in western and central Asia in relation to their dispersal patterns from
cultural perspectives.

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Discussion on the evolutionary status of mainland East Asian late
Middle Pleistocene archaic Homo________________________________________________________________________________

Xing SongKey Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology andPaleoanthropology, Chinese Academy of Sciences________________________________________________________________________________
Late Middle Pleistocene is a crucial period for the origin of H. sapiens. At this time, mainland East
Asia was occupied by a group of archaic Homo, e.g., Dali, Jinniushan, Maba, and Chaoxian. Regarding
for its taxonomy, archaic H. sapiens, H. heidelbergensis, China archaic, archaic human, and late
Middle Pleistocene non-erectus hominin or others have been proposed. However, none of this receive
consistent approvals. Previous studies have revealed that these archaic Homo was characterized by amosaic morphology of primitiveness and progressiveness. This state could be observed in cranial,mandibular, dental, and even postcranial materials without specificity of anatomical structures. In
chronological age, more evidences support an approximately contemporaneity among archaic Homofrom East Asia (~300-105 ka), Neanderthals from Europe (~250-40 ka), and Denisovans from Siberia
and the edge of Tibet Plateau (~287-55 ka). Molecular evidences showed that Neanderthals andDenisovans belong to sister group, and they could hybridize to leave offspring as well as contribute to
the ancestry of modern humans. Related to the Neanderthals and Denisovans, the taxonomic status of
East Asian archaic Homo and its phylogenetic relationships with these two groups are unclear.
According to the studies on the available materials, the classic features that are diagnostic of
Neanderthals are rare in East Asian archaic Homo and there is no solid evidence to lump the East Asianarchaic Homo into the branch of Neanderthals. Another possibility about the evolutionary status of
East Asian archaic Homo is that it belongs to Denisovan. The findings of Xiahe mandible and its
Denisovan classification increase this possibility. In addition, the reconstruction of Denisovan
anatomy using DNA methylation maps match this special population with that of Xuchang, anotherarchaic Homo from East Asia. However, the morphological comparisons between East Asian archaic
Homo and Denisovans reveal variability between them, and it decreases the possibility that theavailable East Asian archaic Homo represent Denisovans. Nevertheless, morphological studies on the
cranial, mandibular, and dental materials have revealed an increasing trend of diversity, the possibility
that more Denisovans will be found in mainland East Asia in the future cannot be ruled out. Taking
into considerations of available morphological and/or molecular evidences, East Asian archaic Homo,
Neanderthals, and Denisovans living at approximately the same time but in separated geographic
territories could represent three separated evolutionary branch rooted in mid-Middle Pleistocene
hominins.

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Denisovans from Baishiya Karst Cave on the Tibetan Plateau________________________________________________________________________________

Dongju Zhang1, Frido Welker2, Chuan-Chou Shen3, Bo Li4, Jean-Jacques Hublin5,
Svante Pääbo6, Qiaomei Fu7, and Fahu Chen8
1)

College of Earth and Environmental Sciences, Lanzhou University

2)

Department of Biology, University of Copenhagen

3)

Department of Geosciences, National Taiwan University

4)

Centre for Archaeological Science, School of Earth, Atmospheric and Life Sciences, University of Wollongong

5)

Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology

6)

Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology

7)

Institute of Vertebrate Paleontology and Paleoanthropology

8)Institute of Tibetan Plateau Research, Chinese Academy of Sciences________________________________________________________________________________In East Asia, mysterious Denisovans so far are only found in Baishiya Karst Cave (BKC) on the
Tibetan Plateau. A human mandible found in the 1980’s in BKC is confirmed to be from a Denisovan
at least 160-thousand-years-old based on paleoproteomic analysis and U-series dating of carbonate
crust outside of the fossil, providing the first Denisovan fossil evidence outside of Denisova Cave andthe earliest human occupation evidence on the Tibetan Plateau. Subsequent archaeological excavation
in BKC reveals that prehistoric human occupied the cave for a long time from the late Middle
Pleistocene to the Late Pleistocene (Chen et al., 2019). Comprehensive studies of stratigraphy,
chronology, archaeology and mitochondrial DNA extracted from the sediments in BKC suggest that
Denisovans occupied the cave ~100 ka and ~60 ka and possibly as recently as ~45 ka (Zhang et al.,2020). Analysis of the rich lithic and faunal remains collected during the excavation shows that simple
core and flake technology was mainly used for lithic production and various wild animals, including
rhinos (probably wooly rhinos) and hyena, were hunted. Paleoenvironment studies in the site andGanjia Basin where BKC is located indicate that Denisovans had experienced large environment
fluctuations in this high-altitude region. The long-term and intensive occupation of BKC by
Denisovans suggests that they may have adapted to life at high altitudes and may have contributed
such adaptations to modern humans on the Tibetan Plateau.
References
Chen, F., Welker, F., Shen, C.-C., Bailey, S.E., Bergmann, I., Davis, S., Xia, H., Wang, H., Fischer, R., Freidline, S.E., Yu, T.-L.,
Skinner, M.M., Stelzer, S., Dong, G., Fu, Q., Dong, G., Wang, J., Zhang, D., Hublin, J.-J., 2019. A late Middle Pleistocene
Denisovan mandible from the Tibetan Plateau. Nature 569, 409-412.
Zhang, D., Xia, H., Chen, F., Li, B., Slon, V., Cheng, T., Yang, R., Jacobs, Z., Dai, Q., Massilani, D., Shen, X., Wang, J., Feng, X.,
Cao, P., Yang, M.A., Yao, J., Yang, J., Madsen, D.B., Han, Y., Ping, W., Liu, F., Perreault, C., Chen, X., Meyer, M., Kelso, J.,
Pääbo, S., Fu, Q., 2020. Denisovan DNA in Late Pleistocene sediments from Baishiya Karst Cave on the Tibetan Plateau. Science
370, 584-587.

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The complexity of the Middle Paleolithic industries in China andimplications for trajectories of human evolution in the region________________________________________________________________________________

Xing GaoThe Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences________________________________________________________________________________The Middle Paleolithic is a controversial issue in Paleolithic research in China. While someresearchers argue that there is no Middle Paleolithic industry in China comparable to that of thewestern Eurasia, others argue that there are some recognizable Middle Paleolithic cultural elements inthe archaeological remains accumulated in China. Such arguments reflect the reality that technologicaldevelopment in China during the period coinciding with the “Middle Paleolithic” of western Eurasia
is multifaceted and complicated, and deserves in-depth investigation in order to comprehend its factual
nature and the dynamics behind it. Based on current discoveries, Paleolithic industries in that periodof time in China exhibit at least three developmental trends, that is, the traditional core-flake industries(i.e. the small flake tool tradition in North China and the pebble tool tradition in South China), theAcheulean-like tradition, and the Mousterian-style techno-complexes. While the first trend prevailed
all the way from the Lower Paleolithic to the Upper Paleolithic, the latter two seem to be only
temporary and regional cultural phenomena. Why the core-flake tradition succeeded so long and sostrong, and where were the source of the Acheulean-like and the Mousterian-style techno-complexes?
These are hotly debated questions that need to be studied further. In addition, more attention shouldbe paid to probe into a fundamental issue, that is, the creator of those complex cultural remains. We
need to examine the origin and migration of certain hominid groups behind such cultural variables, aswell as their social constrain and adaptation strategies.

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What we know about Denisovans: An overview of current fossils,archaeology, chronology and geographic spread________________________________________________________________________________

Katerina Douka1,2,3
1) Department of Evolutionary Anthropology, Faculty of Life Sciences University of Vienna
2) Research Associate, Max Planck Institute for the Science of Human History
3) School of Archaeology, University of Oxford________________________________________________________________________________
The first-ever identified Denisovan was discovered at the eponymous site, Denisova Cave, in the
Siberian Altai over a decade ago. It derived from a relatively high part of the stratigraphic sequence,
and while it was initially reported to be as young as 30 ka, it was indirectly dated to around 60 ka(Douka et al., 2019; Jacobs et al., 2019). More Denisovans were discovered from the site over theyears, both through excavation as well as through biomolecular methodologies, in particular, screeningof thousands of bones using palaeoproteomics and of sediment using aDNA techniques. Yet, until
recently little was known of the first appearance of Denisovans at the cave and in the region moregeneral, and similarly of their wider geographic distribution, their subsistence strategies, interactions
with other hominin species, and eventual demise.
In this paper, I will summarise what we know about the Denisovans, mostly through the lens of the
Denisova Cave findings. Recently, we discovered three hominin bones from the earliest archaeologicallayers of Denisova’s East Chamber (layer 15 and 14). Genetic analyses shown that they carrymitochondrial DNA of the Denisovan type (Brown et al., 2022), placing therefore Denisovans as thefirst hominin occupants of the site. Indirectly dated to ~200 ka they fall at MIS 7, a very warm
interstadial that would have favoured fauna and plant resources to thrive locally and would have render
the Altai mountains a favourable spot for human occupation. The stratigraphic context of these early
Denisovans contains a wealth of archaeological material in the form of lithics and faunal remains,
allowing us to determine the material culture associated with these early hominins and explore their
behavioural and environmental adaptations.
Finally, I will attempt to place Denisovans in their broader Eurasian context by discussing how the
new hominin remains, their chronological position and associated archaeological evidence compare
to those of other locations in Asia where Denisovans are assumed to have lived.
Acknowledgments
This work received funding from the ERC under the European Union’s Horizon 2020 Research and Innovation Programme grant
agreement no. 715069 (FINDER).

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Douka, K., Slon, V., Jacobs, J., Bronk Ramsey, C., Shunkov, M.V., Derevianko, A.P., Mafessoni, F., Kozlikin, M.B. Li, Bo, Grün, R.,
Comeskey, D. Devièse, T., Brown, S., Viola, B., Kinsley, L., Buckley, M., Meyer, M., Roberts, R.G., Pääbo, S., Kelso, J., Higham,
T. 2019. Age estimates for hominin fossils and the onset of the Upper Palaeolithic at Denisova Cave. Nature 565(7741), 640-644.
Jacobs, Z., Li, B., Shunkov, M.V. et al. 2019. Timing of archaic hominin occupation of Denisova Cave in southern Siberia. Nature
565, 594–599.
Brown, S., Massilani, D., Kozlikin, M.B, Douka, K. 2022. The earliest Denisovans and their cultural adaptation. Nature Ecology and
Evolution 6, 28-35.

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Palaeoproteomics for human evolution________________________________________________________________________________

Frido Welker
Section for Ecology and Evolution, Globe Institute, University of Copenhagen________________________________________________________________________________The elucidation of the complex evolutionary relationships between Neanderthals, Denisovans, andmodern humans has recently revealed a new picture of our own ancestral past. How preceding hominin
populations fit into this framework is partly unknown. It has now become clear that Pleistocene
hominin populations were highly diverse, including several newly described hominin populations.
These hominins were present across Africa and Eurasia, with large portions of the hominin fossil
record outside the reach of ancient DNA research. Palaeoproteomic analysis of skeletal proteomes has
recently emerged as a potential additional biomolecular approach across the Pleistocene that reaches
much further back in time compared to ancient DNA. As a result, palaeoproteomics could provide
molecular evidence on hominin evolutionary relationships on a global scale not reachable via othermolecular methods. Achieving this aim requires overcoming existing limitations on sample size and,simultaneously, optimization of the information content recovered from ancient skeletal proteomes.
Based on several case studies and ongoing research of the ERC Project PROSPER, I will discuss andpresent recent developments in the field of palaeoproteomics that might make it possible to retrieve
sufficient quantities of proteomic sequence information from Middle Pleistocene hominins forevolutionary analysis.

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Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology, Palaeoanthropology, and Genetics

Session 2
Origin and expansion of
Homo sapiens in western Eurasia

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Keynote Lecture

Origin and expansion of Homo sapiens in western Eurasia______________________________________________________________________
Jean-Jacques Hublin1,2
1) Collège de France2) Max-Planck Institute for Evolutionary Anthropology______________________________________________________________________
Genetic data from extant humans and the fossil record indicate that our species originated in Africa.
The first Homo sapiens are documented in Northwest Africa 300,000 years ago, in association withthe oldest Middle Stone Age assemblages. However, it is likely that different African populations
contributed to the appearance of the later, so-called "modern" forms of Homo sapiens. During thelate Middle and early Late Pleistocene, “Green Sahara” climatic episodes facilitated the spread of
African populations towards southwest Asia. Their presence in the Near-East is documented in the
paleontological record from 190,000 years BP. Contact with Eurasian Neandertals may haveoccurred even earlier, as suggested by the introgression of Y-chromosome and mitochondrial DNA
haplotypes of African origin into the genome of western Eurasian Neandertals. In contrast, thesubsequent expansion of our species into the mid-latitudes of Eurasia seems to have little to do withenvironmental conditions, and the earliest presence of modern Homo sapiens in Eastern Europe over
45,000 years ago occurred in a cold climate, as evidenced by Bacho-Kiro Cave (Bulgaria). Most
likely, the spread of the initial Upper Paleolithic assemblages associated with these pioneer groups
from Central Europe in the west to the Altai and Mongolia in the east was triggered by cultural anddemographic factors. This first wave of modern settlement was not entirely successful in Europe.
The first modern European populations did not leave detectable traces in the genome of present-day
Europeans, and Neandertals persisted in far western Europe at least until 40,000 years ago. During
this long period of coexistence on a continental scale, complex biological and cultural interactions
took place. They still remain largely to be deciphered. Only with the later arrival of the makers of
the Aurignacian complex did the complete replacement of the local archaic populations take place.

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Pre-Neanderthals and modern humans in the Levant during the late
Middle Pleistocene: A perspective from the Nesher Ramla Homo fossil________________________________________________________________________________

Israel Hershkovitz1,2
1)

The Dan David Center for Human Evolution and Biohistory Research, Tel Aviv University

2)

Department of Anatomy and Anthropology, the Shmunis Family Anthropology Institute, Sackler Faculty of
Medicine, Tel Aviv University________________________________________________________________________________The recently published late Middle Pleistocene Homo fossil found at Nesher Ramla, Israel, hasreignited debate on the nature of the local fossils during this period, their cultural and biologicalrelationships as well as their association with Middle Pleistocene Eurasian populations. In this talk I
will describe the fossil, present the results of the comparative study, and discuss them within a wider
late Middle and early Late Pleistocene anthropological record of Eurasia.

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Multiple pathways to the Upper Paleolithic? Geographic variability in
the Middle to Upper Paleolithic transition in the Levant______________________________________________________________________
Seiji Kadowaki
Nagoya University Museum, Nagoya University______________________________________________________________________A traditional explanation for the Middle to Upper Paleolithic (MP-UP) transition in the Levant is theintroduction of new cultural/behavioral patterns associated with incoming anatomically modern
humans (AMHs) that replaced Neanderthals. Recently, some researchers have recognized both
continuous and discontinuous cultural elements from the Late MP to the Initial Upper Paleolithic (IUP)
and suggested autochthonous cultural development within indigenous populations with someinfluence of incoming groups. The latter view is congruent with an idea of coexistence of Neanderthals
and AMHs in the Levant during the MP that led to the formation of polymorphic populations throughinteraction and interbreeding. The intensification of such social interactions, instead of a population
turnover, has recently been proposed as a driver for the MP-UP cultural changes.
To discuss the above issue from a new perspective, this paper presents archaeological records thatsuggest geographic variability in the MP-UP cultural changes in the Levant. I review chronologicaland lithic technological data of several IUP and Early Upper Paleolithic (Ahmarian) assemblages,
including my original data from the fieldwork in southern Jordan and northern Syria. I suggest thatgeographically variable technological trajectories from the IUP to the Ahmarian can serve as robust
evidence that characterizes the MP-UP transition in the Levant. This is because the technologicaltrajectories revealed by stratigraphic and geographic evidence are less vulnerable to chronologicaluncertainties arising from the difficulty in accurate dating for the period in question. Thus, even if thecurrent chronology for the Late MP, IUP or the Ahmarian are to be changed or refined in future, theregionally different cultural trajectories from the IUP to the Ahmarian will remain stable evidence forthe geographically variable pattern of cultural changes that provide important implications for humanbiogeographic phenomena in the Levant and our understanding in how the MP-UP cultural transition
took place.

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MIS3 palaeoclimate and palaeoenvironment in western Eurasia______________________________________________________________________
Andrea Columbu
Department of Earth Sciences, University of Pisa______________________________________________________________________
Earth’s climate is the end-product of a multitude of interactions and energy exchanges between several
different subsystems, comprising atmosphere, hydrosphere, land surface, cryosphere and biosphere.
All components are intimately linked or coupled with each other, such that changes in one subsystem
may involve compensatory changes throughout the entire climate system. Thus, climate concurrently
controls, and is controlled by, the Earth’s surface environment where Humans live in.
The Arctic Region and the Atlantic Ocean are the “climate-regulators” of the Northern Hemisphere(and possibly the entire Planet), meaning that major changes occurring in these two broad regions may
have effects on in-land climate and environments. However, how major climate changes
propagate/propagated and impact/impacted throughout the vast Eurasian landmass is/was dependent
upon: i) the magnitude and characteristics of climate changes; and ii) the geographical peculiarities ofa certain area within the vast landmass. This means that specific hemispheric (palaeo)climate changes
must be evaluated at specific locations. Palaeoclimate/environment geological archives such as
speleothems, lacustrine and marine sediments, tree rings, etc, are highly useful to reconstruct ancient
climates at specific locations within the framework of hemispherical-to-global climate variations.
From a general perspective, Marine Isotopic Stage 3 (MIS3, ~57 to ~29 kiloyears BP) is a transitional
climate stage placed between the last interglacial (MIS 5e) and the last glacial maximum (MIS1).
MIS3 is characterised by important intra-stage climate fluctuations in the Arctic Region and theAtlantic Ocean, which in turn affected continents. In this regard, the aim of this talk is to explore theeffects of MIS3 hemispheric-to-global climate instability throughout the western Eurasian territory,
by examining palaeoclimate/environmental archives from selected locations.

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The genomic profile of Upper Paleolithic European hunter-gatherers_____________________________________________________________________________

Cosimo Posth
Archaeo- and Palaeogenetics group, Institute for Archaeological Sciences & Senckenberg Centre for Human Evolution
and Palaeoenvironment, University of Tübingen_____________________________________________________________________________
After reaching Europe, modern humans relied on a foraging lifestyle for nearly 40,000 years and thereconstruction of their demographic history along this time span is of great interest. Paleogenetic
analyses of Upper Paleolithic hunter-gatherers have contributed to the understanding of population
structure and genetic turnovers in Ice Age Europe. Through temporal and geographic transects ofgenomic data it is in fact possible to infer past population movements and to identify their potential
links with the climatic and archaeological records. However, our knowledge of the genetic history ofEuropean hunter-gatherers is still limited due to the relatively sparse and often poorly preserved humanskeletal remains from that period. To date, genetically analyzed individuals older than 40,000 years
ago are found to represent deeply divergent out-of-African lineages that did not leave traces in thegene-pool of present-day Europeans. On the contrary, individuals dating after 37,000 years ago belong
to at least two main lineages that are primarily nested within the European branch after its split from
Asians. Those ancestries survived in different European areas until the beginning of the Last Glacial
Maximum (LGM, ~25-19,000 years ago), the coldest phase of the last Ice Age. The role of proposed
climatic refugia for human populations during the LGM is discussed here, as well as the re-peoplingof Europe after this event. Particularly important is the appearance of a new lineage in southern Europe,
found to carry genetic ties with Near Eastern populations. This lineage further spread across Europe
after 14,000 years ago, in conjunction with a major warming event, and largely replaced the preceding
genetic landscape. However, not all European regions were equally influenced by this demographic
shift. In the Iberian Peninsula the genetic turnover was minor and in eastern Europe a geneticallydistinct population emerged after the LGM. By expanding the distribution of hunter-gatherer genomic
data through time and space it is thus becoming possible to delineate at a higher resolution the genomictransformations and interactions that took place in Europe during the Upper Paleolithic.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

The Initial Upper Palaeolithic at Bacho Kiro Cave and in the Eastern
Balkan area________________________________________________________________________________

Tsenka Tsanova
Palaeoanthropology, Senckenberg Centre for Human Evolution and Palaeoenvironment, Institute for Archaeological
Sciences, Eberhard Karls University of Tübingen________________________________________________________________________________

The cave is located in the eastern part of the Balkans Peninsula, in northern Bulgaria, at thegeographical lobby of the migration routes along the Danube corridor and has been reinvestigatedsince 2015. The previous excavations from the 1970s had revealed a sequence, spanning from theMiddle to the Upper Palaeolithic with rich archaeological assemblages, known as Bachokirian from
Layers 11 and 11a (Kozłowski 1982). The new excavation uncovered similar stratigraphy to thosepreviously published. From techno-typological perspective, the presence of Levallois features
combined with UP tool types allows to reattribute the Bachokirian to the Initial Upper Palaeolithic(IUP) (Kuhn and Zwyns 2014). The new human remains mostly detected by ZooMS are radiocarbondated to ca 45 ka cal BP (Hublin et al. 2020; Fewlass et al. 2020) and represent the largest number ofrecovered individuals in the context of their stone and bone tools, ornaments, and rich fauna remains,at this period of time.
The Layer I is in a good state of preservation, no evidence of mixing of separate entities, and theassemblage is characterised by: 1) artefacts made on allochthonous Aptian and Campanian flint; 2)
off-site production and transportation of finished products (blades); 3) blade technology withmorphology close to Levallois: large, straight blades with thick platforms, detached by direct
percussion with hard hammer; 4) a high proportion of burnt artefacts and weathered surfaces; 5) a highrate of retouched tools on-site, morphological and typological variability among pointed blades andtools manufactured, with diagnostic impact fractures on some pointed blades; 6) high fragmentation
rate, reworking and reshaping of tools; 7) intense reduction of the artefacts by bipolar knapping,sometimes related to their use as wedges for working hard organic material (Horta et al. 2020); 8)
bladelets and small flakes obtained from core-on-flake and bipolar on-anvil percussion, presence ofvarious splintered tools and blanks.
There are other assemblages in the same regional chrono-stratigraphic record, technologically andchronologically compatible with the IUP from Bacho Kiro Layers I and J, which are: Temnata (Layers
VI and 4), Kozarnika (Layer 6/7 id. Level VIII), Samuilitsa II caves and possibly in Toplitsa cave
(Layer 5) (Tsanova et al. 2021). Regional assemblages with Levallois and pointed blades fromexcavations before 1960 (Devetashkata, Tabashkata, Vasil Levski Caves) need to be re-assessed withpossible links to IUP or Levallois-Mousterian traditions (Tsanova et al., in prep.). Finally, the IUP
assemblages lasted after 39 ka cal BP in Temnata cave, Layer 4 (Tsanova, in press).

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References
Kozłowski, J. K. 1982. Excavation in the Bacho Kiro cave, Bulgaria. Final report. In Kozlowski, J. K., (ed.) Warszawa, Państwowe
Wydawnictwo Naukowe, p. 172.
Kuhn, S.L. and Zwyns, N. 2014. Rethinking the initial Upper Paleolithic. Quaternary International 347, 29-38.
Hublin, J.-J., Sirakov, N., Aldeias, V., Bailey, S., Delvigne, V., Endarova, E., Fewlass, H., Hajdinjak, M., Krumov, I., Marreiros, J.,
Martisius, N., Sinet-Mathiot, V., Meyer, M., Popov, V., Sirakova, S., Smith, G., Spasov, R., Režek, Ž., Talamo, S., Welker, F.,
Wilcke, A., Nikolay Zahariev, McPherron, S. P., and Tsanova, T. 2020. Initial Upper Palaeolithic Homo sapiens remains from
Bacho Kiro Cave (Bulgaria). Nature, 581(7808), 1-4.
Fewlass, H., Talamo, S., Wacker, L., Kromer, B., Tuna, T., Fagault, Y., Bard, E., McPherron, S. P., Aldeias, V., Maria, R., Martisius,
N. L., Paskulin, L., Rezek, Z., Sinet-Mathiot, V., Sirakova, S., Smith, G. M., Spasov, R., Welker, F., Sirakov, N., Tsanova, T.,
and Hublin, J.-J. 2020. A 14C chronology for the Middle to Upper Palaeolithic transition at Bacho Kiro Cave, Bulgaria. Nature
Ecology & Evolution, 4(6), 794-801.
Horta, P., Tsanova, T., Sirakov, N., Marreiros, J., Zahariev, N., Sirakova, S., Hublin, J.-J. 2020. Intensive re-use of stone tools as
bipolar implements in the Initial Upper Paleolithic of Bacho Kiro Cave. Poster presented at the 10th Annual meeting of theEuropean Society for the study of Human Evolution, Virtual meeting.
Tsanova, T., Veres, D., Hambach, U., Spasov, R., Dimitrova, I., Popov, P., Talamo, S., Sirakova, S., 2021. Upper Palaeolithic layers
and Campanian Ignimbrite/Y-5 tephra in Toplitsa cave, Northern Bulgaria. Journal of Archaeological Science: Reports, 37,
102912Tsanova, T., (in print). Blades and bladelets technologies at the onset of the Upper Palaeolithic in Eastern Balkans. Preliminary
comparison and chronology of the lithic assemblages from Bacho Kiro, Temnata and Kozarnika caves in North Bulgaria. In
Ruiz-Redondo, A., and Davies, W. (eds.) The Prehistoric Hunter-gatherers of South-eastern Europe: recent research,
Cambridge, Proceedings of the British Academy.

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Cultural perspectives on early dispersals of Homo sapiens in Central
Europe and Eastern European Plains________________________________________________________________________________

Andrea Picin
Bereich für Ur- und Frühgeschichtliche Archäologie, Friedrich Schiller Universität Jena________________________________________________________________________________
During the Initial and Early Upper Paleolithic, Levantine groups of Homo sapiens reached newgeographic distributions moving northwards into unfamiliar European environments. In thesedispersals along the Danube River and the Mediterranean coast, Homo sapiens was geared with newtechnologies and subsistence strategies that allowed the fully exploitation of colder ecological habitats.
Thus far, researches in Central Europe support the hypothesis that these new settlements wererestricted to certain latitudes and the territories above 49° north were ecological barriers for the newgroups due to the harsh climates and the high seasonality of the biotic resources. Furthermore, even if
the environmental conditions remained similar during the second half of MIS 3, it is thought that only
after 35 ka BP, Homo sapiens was able to cross these ecological impediments and spread northwards(e.g. north of the Carpathians). This paper aims to review the current archaeological evidence fromCentral Europe and the Eastern European Plains, and present new data from Poland, the territory
halfway between the two areas. While the Initial Upper Paleolithic signals continue to be weak in theregion, the re-examination of several lithic assemblages support the view of cyclical visits since the
Early Aurignacian.

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ERC project SUCCESS: The earliest migration of Homo sapiens insouthern Europe________________________________________________________________________________

Stefano Benazzi
Department of Cultural Heritage, University of Bologna________________________________________________________________________________The time period between 50,000-40,000 years ago (ka) is a crucial event in human evolution, as H.
sapiens expanded out of tropical areas into Eurasia replacing or partially absorbing local archaic
humans, among which were the Neanderthals. To date, the time and mode of this major global
replacement of populations remain unknown. Within this context, new evidence portrays Italy as a
keystone region in answering questions surrounding this transition due to its geographic position,
ecological variability, and key archaeological Middle-to-Upper Paleolithic sites, yet Italy has been
largely absent in research. In this talk, I’m going to present the results of an ERC Consolidator Grant(n. 724046 - SUCCESS) that tackles this issue. The SUCCESS project is organized in 5 work packages(i.e., Fieldwork, Paleoecology, Analysis of faunal and human remains, Study of cultural objects,
Analysis and modeling of cultural change), with a Research Team composed of seven postdoctoral
fellowships and three PhD students. During the 5-year project, we deepened our understanding of the
Italian paleoclimatic/paleoenvironmental variability in terrestrial ecosystems between 50-40 ka at subcentennial scale, and we studied new human remains retrieved from Italian deposits dated to theMiddle-to-Upper Paleolithic period. Moreover, we provided the most extensive study of the Uluzzian
technocomplex, discarding its relationship with the Mousterian culture while supporting its inclusion
within the early Upper Paleolithic. Among others, we determined that the backed pieces were used as
hunting armatures on projectiles used with a spearthrower or a bow; moreover, we also found that theshell ornaments retrieved from the Uluzzian levels of Grotta del Cavallo represent the earliest known
shell ornament-making context in Europe.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Advanced projectile technology of the earliest Homo sapiens in Europe________________________________________________________________________________

Katsuhiro Sano1, Simona Arrighi2, 3, Lisa Vaccari4, Stefano Benazzi2 and Adriana
Moroni3
1)

Center for Northeast Asian Studies, Tohoku University

2)

Department of Cultural Heritage, University of Bologna

3)

Department of Environment, Earth and Physical Sciences, University of Siena

4)

Elettra-Sincrotrone Trieste S.C.p.A.

________________________________________________________________________________
Apparently, Neanderthals and Homo sapiens coexisted in Europe for over 5,000 years. However, little
is known about why Homo sapiens could increase their population size after migrating to Europe andsuccessfully occupy new territories, while autochthonous Neanderthals went extinct ~40,000 years
ago. To better understand the replacement of Neanderthals by Homo sapiens, we studied modern
human hunting weapons, directly relating to their subsistence strategies. We examined 146 crescentshaped backed pieces retrieved from the Uluzzian culture of Grotta del Cavallo (Southern Italy) dated
between 45 - 40 ka. The backed pieces were macroscopically and microscopically analyzed usinga Hirox digital microscope and results were compared with use-wear patterns on experimental
samples. Through this analysis, diagnostic impact fractures and microscopic impact linear traces
were found on numerous backed pieces, demonstrating that they were used as hunting weapons.
The diagnostic impact fractures showed the similar patterns of experimental samples delivered by aspearthrower and a bow, but significantly different from those observed on throwing and thrusting
samples. Recent use-wear analysis of Initial Upper Palaeolithic points from the Boker Tacktit in theLevant also indicates that they were projected with high impact velocity, suggesting that Initial Upper
Palaeolithic points may have performed the same function in Europe as well. In contrast, no Middle
Palaeolithic points used by Neanderthals show similar impact patterns and they are overall much larger
than Upper Palaeolithic points. These results signify that Homo sapiens migrating into Europe
equipped themselves with mechanically delivered projectile weapons, such as a spearthrower-darts ora bow-and-arrows, which had higher impact energy. As the advanced hunting strategy isstraightforwardly related to a competitive advantage, this study offered important insight to understandthe reasons for the replacement of Neanderthals by Homo sapiens.

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What a precise clock! ERC project RESOLUTION provides more refinechronology of the Initial and Early Upper Palaeolithic in Europe________________________________________________________________________________

Sahra Talamo
Department of Chemistry G. Ciamician, Alma Mater Studiorum, University of Bologna________________________________________________________________________________
Radiocarbon dating is the most widely used dating method for archaeologists. However, the 14C datedoes not correspond to the true calendar age, but to obtain it, the varying level of 14C back throughtime needs to be reconstructed. Briefly, we need a calibration curve (Reimer et al., 2020).
Radiocarbon calibration for the past 14,000 years is highly precise and of high resolution (annual to a
decade) due to tree-ring chronologies, but it is still rather coarse in the Initial and Early Upper
Palaeolithic, where the carbon archives are connected only indirectly to the atmosphere (speleothems,
coral reefs, marine and lake sediments). However, the resolution of the most used atomic ‘clock’ in
the Glacial will be improved by new findings of Glacial trees in New Zealand (Kauri) and in the
Mediterranean (Italy, Portugal), linked to the ice-core timescale via another cosmogenic isotope,
Beryllium-10 (Adolphi et al., 2017).
Using these tree-ring-based data, we can begin to calibrate more accurately the 14C ages associatedwith breakthrough discoveries of human evolution. The most recent discovery showed that Homosapiens was already in Europe around 46,000 years ago at Bacho Kiro in Bulgaria (Hublin et al., 2020;
Fewlass et al., 2020), propagating east-west along the Mediterranean rim within a fairly short time andreaching the westernmost part of Europe between 41,000 and 38,000 years ago (Haws et al., 2020).
Moreover, the question about the beginning of the artistic development by Homo sapiens is stillchallenging because of the limited chronological information. The recent discovery of a pendant in
Poland, dated 41,500 years ago, plays a unique role in demonstrating the importance of directly dating
objects of Palaeolithic art to understand the origin of communication, celebration, and expression of
Homo sapiens in Europe (Talamo et al., 2021).
These findings suggest that the history of our evolution in Europe has been particularly complex,
leading us to emphasize the importance of having a precise and detailed ‘clock’ that can determine theaccuracy of the time intervals that mark the course of our history.
Acknowledgements
S. Talamo has received funding from:
1) European Research Council under the European Union’s Horizon 2020 Research and Innovation Programme (grant agreement No.
803147- RESOLUTION, https://site.unibo.it/resolution-erc/en).
2) PRIN 2020, the Research Projects of Major National Interest (Grant agreement Code 20209LLK8S - DYNASTY)

References

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Reimer PJ, et al. 2020. The IntCal20 Northern hemisphere radiocarbon age calibration curve (0–55 cal kbp). Radiocarbon 62(4):725757.
Adolphi F, et al. 2017. Radiocarbon calibration uncertainties during the last deglaciation: Insights from new floating tree-ringchronologies. Quaternary Science Reviews 170:98-108.
Hublin J-J, et al. 2020. Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria. Nature 581(7808):299-302.
Fewlass H, et al. 2020. A 14C chronology for the Middle to Upper Palaeolithic transition at Bacho Kiro Cave, Bulgaria. Nature
Ecology & Evolution 4(6):794-801.
Haws JA, et al. 2020. The early Aurignacian dispersal of modern humans into westernmost Eurasia. Proceedings of the National
Academy of Sciences 117(41):25414-25422.
Talamo S, et al. 2021. A 41,500 year-old decorated ivory pendant from Stajnia Cave (Poland). Scientific Reports 11(1):22078

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INTERNATIONAL SYMPOSIUM
Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology, Palaeoanthropology, and Genetics

Special Session
The Japanese Palaeolithic:
Isolation and cultural exchanges

INTERNATIONAL SYMPOSIUM

Special Lecture

Another Palaeolithic frontier: Modeling the earliest seafaring in East
Asia________________________________________________________________________________

Yousuke Kaifu
The University Museum, The University of Tokyo________________________________________________________________________________The rise of voyaging technology beyond nearshore boating was a key for early modern humans toexponentially expand their habitable territory on the globe. However, apart from intensive discussion
on the earliest such evidence from Wallacea, the developmental process and regional variation of theLate Pleistocene seafaring remain unclear. To contribute this issue, I present a synthetic model for
Palaeolithic seafaring in another region of the western Pacific, the Ryukyu Islands (Ryukyus),
southwestern Japan. Here, some of the islands were more than 100 km away and invisible beyond thehorizon, and one of the world’s strongest ocean currents, Kuroshio, intervened the seaways. Despite
these challenging situations, Palaeolithic sites appeared throughout much of the 1,200 km chain of theislands ~35,000–30,000 years ago. By integrating currently available information from archaeology,
skeletal morphology, genetics, palaeogeography, oceanography, and our own experimental projectcalled ‘Holistic Reenactment Project of Voyages 30,000 Years Ago’,(https://www.kahaku.go.jp/research/activities/special/koukai/en/), I discuss probable migration routes, possible

watercrafts, preparation and strategy needed for successful maritime migrations in the region, andother issues relevant to deeper understanding of the origins and development of human maritime
activities.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

The spatio-temporal patterns of early Upper Palaeolithic assemblages
in the Japanese islands________________________________________________________________________________

Katsuhiro Sano1, Shunsuke Totsuka2, Masami Izuho3, Kazuki Morisaki4
1)

Center for Northeast Asian Studies, Tohoku University

2)

Gradual School of Arts and Letters, Tohoku University

3)

Gradual School of Humanities, Tokyo Metropolitan University

4)

Gradual School of Humanities and Sociology, The University of Tokyo________________________________________________________________________________In the Japanese islands, the number of Palaeolithic sites abruptly increases after c. 38 kcal BP, whichcoincides with the expansion of Homo sapiens into these islands. The lithic assemblages between 38
and 30 kcal BP, assigned to the early Upper Paleolithic (eUP), are characterized by trapezoids, pointed
blades, and edge-ground axes. The Japanese eUP assemblages are quite different from thechronologically simultaneous Upper Palaeolithic assemblages in the neighboring regions, such as
Korea, China, and the Russian Far East. Therefore, the distinctive features of the Japanese eUP may
have been developed when modern human hunter-gatherers adapted into the environment in theJapanese islands. In this study, we examine spatio-temporal patterns of the eUP assemblages in theJapanese islands and discuss when and how the important components of these emerged. Based on theBayesian models of reliable radiocarbon dates, we confirmed that trapezoids first appeared at ~37,500
cal BP, then blade production started at ~36,500 cal BP. Thereafter, both the trapezoid- and bladebased assemblages rapidly increased between 35 and 34 kcal BP, and the important eUP components,
such as edge-ground axes and circular lithic concentrations, are also prevalent at numerous sites.
However, from 34 kcal BP onward, the eUP sites drastically decreased, and the numbers of trapezoids,
edge-ground axes, and circular lithic concentrations are reduced. The concept of blade production has
also changed during this time. While triangular blades were unidirectionally removed from a narrowside of a core without preparing the striking platform in the earlier phase of the eUP, a new blade
reduction concept represented by late Upper Palaeolithic (lUP) assemblages in Japan (blade detachingfrom a wide face of a prismatic or semi-cylindrical core with the preparation of the striking platform)
gradually became predominant after 34 kcal BP. The spatio-temporal patterns of the eUP assemblages
indicate that although the representative features of the eUP prevailed when the population density
was high, the eUP components have declined thereafter according to the reduction of the site density
and instead some lUP characteristics progressively emerged.

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Middle and late Upper Palaeolithic in the Japanese archipelago:
Local development and continental influence________________________________________________________________________________

Kazuki Morisaki
Gradual School of Humanities and Sociology, The University of Tokyo________________________________________________________________________________
Previous studies revealed that the radiocarbon dates of the Japanese Upper Palaeolithic ranged from39,000 to 16,000 cal BP from the latter half of marine isotope stage (MIS) 3 to late MIS2. Reliablegeochronology of several regions of the Palaeo-Honshu Island elucidated a millennial-scale
development of local lithic industries after 30,000 cal BP, which is the global Last Glacial Maximum(LGM). In the northern area of Palaeo-Honshu, blade industries based on local cryptocrystalline shale
were dominantly utilized, whereas various flake and/or small blade industries with varying levels of
quality of local raw materials were prevalent in the southern area of Palaeo-Honshu. The diachronicchange in these lithic industries well corresponded to rapid climatic cooling toward MIS2, which isevidenced by a global change in sea-level and palaeo-vegetation study. The inter-regional
diversification of lithic industries occurred in parallel with the population decline of major species
composing large mammal fauna at this time, suggesting that foragers began exploiting local lowmobility mammals.
As represented by the diffusion of diagnostic stemmed points on blade from the southern Korean
Peninsula, several continental influences on the Palaeo-Honshu during MIS2 were recognized.
Stemmed points on blade were mainly introduced into the present Kyushu, which is the southwesternedge of Palaeo-Honshu, during a very brief period, i.e., to be exact, between 29,300 and 27,500 cal
BP, and rapidly changed their form into flake-based ones. This observation indicated short-term/smallscale human migration between the Korean Peninsula and Kyushu.
The end of the LGM circa 20,000–18,000 cal BP witnessed another continental influence: theintroduction of microblade technology throughout the Palaeo-Honshu Island. Although earlymicroblade technology, for example, prismatic types, may be explained by the technological evolutionof a preceding small blade technological tradition and informational transmission from an adjacentcontinent, the emergence of wedge-shaped microblade technology circa 18,000–16,000 cal BP may
haveaccompanied

human

migration

from

Sakhalin/Hokkaido/Kuril Peninsula.

32

continental

regions

includingthe

Palaeo-INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Ecosystem change and decisions for pottery using foraging:
Late Pleistocene cases from the Japanese archipelago________________________________________________________________________________

Fumie Iizuka
Center for Northeast Asian Studies, Tohoku University________________________________________________________________________________In the Amur River basin and islands of Japan, where the earliest ceramic vessels are unambiguously
associated with the terminal Pleistocene, the advent of pottery occurred in the context of broadening
diet. A case from southern Kyushu of southern Japan furthermore suggests that the first indication ofvessel adoption was by ca. 15,000 cal BP associated with a microblade using mobile foraging in theabsence of megafauna. A substantial adoption of pottery began during the Bølling/Allerød in the
Incipient Jomon, ca. 14,000/13,500-12,800 cal BP. In places such as Tanegashima Island to the southof mainland Kyushu, significantly broadened diet and increased sedentism, and adoption of a varietyof new lithic tools and features are observed. It is inferred that change occurred in the context ofresource abundance in a forested, ecotone environment (influenced by the warm Kuroshio current),and sea level rise disconnecting Tanegashima from the mainland Kyushu. Evidence of likely interisland exchange of pottery and lithics is also found. The onset of the Incipient Jomon, therefore, may
be associated with the transition from the Upper Paleolithic-like to Neolithic-like behavioral shiftinterpreted as the forager response to long-term risk related to sea level rise. In this presentation, I
critically evaluate the associations of biome conditions and change, sea level rise, and the timing of
the adoption of pottery and new technology from distinct regions of the Japanese archipelago. Those
cases are compared with the context from southern Kyushu. This study contributes to understanding
hunter-gatherer decision making processes involving pottery adoption, subsistence change, and new
behavioral organization in response to ecosystem change in the terminal Pleistocene in the easternislands of Eurasia. It may also provide comparative perspectives on pottery adopting, hunting andgathering and subsistence strategies of late Pleistocene foragers who migrated to the Americas.

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Ancient genome analysis of human remains in the Japanese archipelago________________________________________________________________________________

Hiroki Oota
Department of Biological Sciences, Graduate School of Science, The University of Tokyo________________________________________________________________________________
The ancient DNA analysis of human remains excavated from the Japanese archipelago started in the
1990s (Horai et al. 1991; Kurosaki et al. 1993; Oota et al. 1995; Shinoda and Kanai 1999). Firstly,
mtDNA HVRs were sequenced in five Jomon individuals (Horai et al. 1991). Subsequently, nuclear
polymorphic loci (variable number of tandem repeats) were examined (Kurosaki et al. 1993), andpopulation studies of ancient mtDNA had also begun (Oota et al. 1995; Shinoda and Kanai 1999; Oota
et al. 1999; Wang et al. 2000).
However, the first application of NGS to ancient genome sequencing in Japan (Kanzawa-Kiriyama etal. 2017) was 10 years later than the application to the Neanderthal genome. This delay was greatly
influenced by the burial environment surrounding ancient Japanese human bones. The climate of theJapanese archipelago is warm and humid, and the soil is acidic because of the volcanic archipelago.
For these reasons, especially on the Honshu (the largest island of the Japanese archipelago), it isdifficult for human bones buried in the soil to remain. Therefore, very few human remains from thepaleolithic period have been found in Honshu. However, we have been getting the Jomon genome datafrom the Japanese archipelago, recently. A draft whole-genome sequence of a 2,500-year-old Jomon
woman from the Ikawazu shell-mound site in the Atsumi peninsula, Honshu, was subsequently
reported (McColl et al. 2018), and a complete whole-genome sequencing of a Jomon individual fromthe Funadomari site in Rebun Island, Hokkaidō, was successfully achieved (Kanzawa-Kiriyama et al.
2019).
In this talk, I will review the ancient genome researches in Japan, and show our recent achievements.
References
Horai et al. 1991. Phylogenetic affiliation of ancient and contemporary humans inferred from mitochondrial DNA. PhilosophicalTransactions of the Royal Society of London, B: Biological Sciences 333(1268):409-16.
Kurosaki et al. 1993. Individual DNA identification from ancient human remains. American Journal of Human Genetics 53(3):63843.
Oota et al. 1995. A genetic study of 2,000-year-old human remains from Japan using mitochondrial DNA sequences. American Journalof Biological Anthropology, 98(2):133-45.
Shinoda and Kanai 1999. Intracemetery Genetic Analysis at the Nakazuma Jomon Site in Japan by Mitochondrial DNA Sequencing.
Anthropological Science, 107(2): 129–140.
Oota et al. 1999. Molecular Genetic Analysis of Remains of a 2,000-Year-Old Human Population in China—and Its Relevance for theOrigin of the Modern Japanese Population. American Journal of Human Genetics, 64(1):250-8.
Wang et al. 2000. The Origin of the Jingwei Gene and the Complex Modular Structure of Its Parental Gene, Yellow
Emperor, in Drosophila melanogaste. Molecular Biology and Evolution, 17(9): 1396–1400.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Kanzawa-Kiriyama et al. 2017. A partial nuclear genome of the Jomons who lived 3000 years ago in Fukushima, Japan. Journal of
Human Genetics, 62(2): 213–221.
McColl et al. 2018. The prehistoric peopling of Southeast Asia. Science 361(6397): 88–92.
Kanzawa-Kiriyama et al. 2019. Late Jomon male and female genome sequences from the Funadomari site in Hokkaido, Japan.
Anthropological Science, 127(2): 83–108.

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INTERNATIONAL SYMPOSIUM
Insights Into Human History in the Eurasian Stone Age:
Recent Developments in Archaeology, Palaeoanthropology, and Genetics

Session 3
The earliest migration of
Homo sapiens into
Central and Northeast Asia

INTERNATIONAL SYMPOSIUM

Problems and prospects in the study of modern-human dispersal in
Northern Asia________________________________________________________________________________

Ted Goebel
Department of Anthropology, University of Kansas________________________________________________________________________________
In northern Asia, specifically southern Siberia from the Altai Mountains to the Lake Baikal region,
information about the origins of the Upper Paleolithic and dispersal of modern humans has grownexponentially in the past 25 years. The region now boasts a chronology of > 100 radiocarbon ages
from > 50 archaeological occupations that chronicle events from 50,000 to 35,000 calendar years ago,
providing an opportunity to precisely measure the timing of the emergence of the Upper Paleolithic.
High-resolution paleoenvironmental records from Lake Baikal and other sources now permit aconsideration of the transition against the backdrop of climate change during Marine Isotope Stage(MIS) 3. Large-scale excavations have provided samples of artifacts adequate for measuring andinterpreting variability among lithic and osseous artifact assemblages. Those same sites have yieldedwell-preserved remains of fauna, facilitating new studies of subsistence behavior. Nonetheless,problems with the record still plague us. First, three distinct hominin taxa (Neanderthals, Denisovans,
and anatomically modern humans) are known to have inhabited the region during MIS 3, and wecannot yet determine which produced southern Siberia’s initial Upper Paleolithic (IUP). Second, manyof the sites are in colluvial settings and individual radiocarbon chronologies suggest they often
represent palimpsests of numerous occupations spanning thousands of years, so that excavations needto carefully consider geological site-formation processes and spatial distributions of finds to measure
the intactness of the IUP occupations. Third, theoretical assumptions for models explaining the IUP’s
emergence are under-developed and need to better consider how ‘behavioral modernity’ is expressedarchaeologically, and how we might best detect behavioral versus cultural patterning in relation to the
spread of modern-human populations. Fourth, outside of the Altai caves, the preceding Middle
Paleolithic remains poorly documented, so that comparisons between it and the IUP are difficult, if
not impossible, to undertake.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Early adaptations into tundra landscape by modern humans: A casestudy of the Initial Upper Paleolithic in Mongolia and Transbaikal,
Russia____________________________________________________________
Masami Izuho
Gradual School of Humanities, Tokyo Metropolitan University________________________________________________________________________________The Initial Upper Paleolithic (IUP) is archaeological phenomena which archaeologists often refer toas one of the “evolved” Upper Paleolithic behaviors: the most widespread, the earliest (~45 ka),
appearing as highly variable in its composition and duration, and discontinuous across Eurasia. In
order to better understand whether the broad similarities of material culture reflect phylogenetic
connections or broad convergence in its local scale, this paper investigates the correspondence
between the chronological sequence from the Initial to Early Upper Paleolithic (EUP) and theecosystem changes during MIS 3 in Mongolia and Transbaikal, Russia (MO-TB), as a test case.
Having a rapid increase in the number of AMS dates in archaeological sites, archaeologists have madea consensus that chronological sequence of the IUP-EUP in MO-TB likely fall within the range of
45,000-30,000 Cal yr BP. Currently, environmental reconstructions are available and those provide
higher-resolution than other datasets mainly due to lake sediment core analyses. Combining thosearchaeological and environmental data, some recent arguments propose that the modern humanoccupations were possibly placed at the onset of climatic ameliorations: IUP coincides with theinterval between Heinrichs event (H) 5 and H4, and EUP between H4 and H3, respectively. We arestill not sure if this expectation relates to the disappearance of occupations in cold-arid events implyingthat the IUP-EUP foragers were not yet adapted to Tundra landscape in MO-TB, simply due to thesampling bias, or because of archaeological visibility. Following this expectation, this paper will
explore a single research question: did modern humans adapt into Tundra landscape during IUP in
MO-TB?

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Exploring the variability of IUP assemblages using a techno-economic
approach: The example of Tolbor-16, Mongolia________________________________________________________________________________

Nicolas Zwyns
Department of Anthropology, University of California Davis________________________________________________________________________________
Lithic assemblages labelled as Initial Upper Paleolithic (IUP) are often described as a combination of
Middle and Upper Paleolithic typo-technological features. This peculiar composition is usuallyinterpreted as reflecting site function, raw material availability, contact between different
populations/species or more broadly, as a technology in transformation. Aside from a common set ofbasic traits, there is little consensus on what is reminiscent of older stone technologies, what does
represent site-specific behaviors and what is fully derived and UP in nature. It is perhaps why the IUP
is described as highly variable unit while what the meaning of such variations remains unclear. Here
we describe in detail an assemblage from the site of Tolbor-16, in Northern Mongolia, dated to ca. 45
ka cal BP. Aside from a derived pattern of blade production that defines the IUP in the region, we
observe a high rate of fragmentation and a low blank frequency of actual blades relative to the evidence
for active production at the site. Estimates of core/blade ratios obtained are relatively low, suggestingthat material was imported in a late stage of reduction and/or that some of the blades were transported
elsewhere. We observe that a substantial part of the retouched tools is produced on flake blanks,
including types occurring in high frequencies among Middle Paleolithic assemblages. The
morphology of the flakes and the absence of a systematic methods of production (e.g. Levallois,
Discoid) point toward the recycling of product coming from the preparation of blade cores. Variations
observed in core morphology is balanced by the redundant use of a peculiar reduction method.
Comparing core volume between the different core types, we suggest that the variations observed inshape could illustrate a process of reduction (as opposed to distinct, independent pathways). Middle
Paleolithic features are mostly typological or related to the percussion technique. Overall, our results
suggest that the combination of traits observed can be explained in techno-economic terms, and do notnecessarily require higher-level explanations such as strict link with older traditions, or contacts
between species. In the present case, a techno-economic approach is helpful to isolate archaic featuresfrom behaviors that reflect equifinality. It warrants against the use of such traits to support
evolutionary interpretations when the data available (e.g. fauna) is not sufficient to fully understandthe nature of the occupation and the subsistence strategies involved.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

Initial Upper Paleolithic of Southern Siberia and Central Asia:
Continuity and variability________________________________________________________________________________

Evgeny P. Rybin1, Kseniya A. Kolobova1 and Arina M. Khatsenovich1
1) Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences________________________________________________________________________________
Substantial progress has been made over the past decade in our understanding of the timing andvariability of the earliest Upper Paleolithic in the South Siberia and eastern Central Asia. In theseregions the emergence of Middle Paleolithic industries, in the technological set of which Levallois
point-and-blade technology played an important role, can be traced from the late MIS-4 stage. Theirchronostratigraphic position at the top of Middle Paleolithic record immediately precedes the local
Upper Paleolithic. In chronological interval 49,000 - 45,000 BP in archaeological complex OH65/UP2 of Kara-Bom site (Russian Altai) is evidenced the existence of complete technological,
typological and symbolic package related to trans-Eurasian techno-complex of Initial Upper
Paleolithic (hereafter IUP). Assemblages of IUP are distributed over an exceptionally vast area of themountain belt of Southern Siberia and Central Asia. The extreme points of their distribution, marked
by stratified sites, are within N 54° - N 38° and E 84° - E 109°. The question of the variability of IUP
assemblages from different regions of its distribution remains open along with the important
discussion about diachronic patterns in lithic technology throughout Middle Paleolithic/Initial Upper
Paleolithic boundaries. A set of statistical methods (NMDS scaling, PCA analysis, LDA analysis) will
be used to achieve this goal. Samples will be grouped in order to determine the variability of theassemblages from the different regions of IUP geographic core according to cultural-chronologicaland territorial grouping. The relative typological and technological homogeneity of the Siberian andCentral Asian IUP in comparison with the Early Upper Paleolithic, substantial difference between IUP
and Middle Paleolithic and narrow chronological difference between the extreme points of IUP
distribution makes possible to suppose the exogenous origin and very fast dispersal of that cultural
event in the middle of Eurasia.
Acknowledgements
Research was supported by Russian Scientific Foundation project # 19-18-00198.

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Understanding the Initial Upper Paleolithic of China in a regional
context of Eastern Eurasia________________________________________________________________________________

Feng Li
School of Archaeology and Museology, Peking University, China________________________________________________________________________________
Initial Upper Paleolithic (IUP) has attracted lots of attentions of scholars who are interested in thetransitional period from Middle Paleolithic to Upper Paleolithic in Eurasia, especially in the easternpart of the landmass recently. As one of the regions, Northern China has played an important role indiscussions on the chronology, diffusion, and adaptation of the IUP assemblages due to the abundantfindings at Shuidonggou locality 1 and other sites. Attempts to understand the IUP of northern China
in a regional context have been practiced, however, there are still much unclear informationconsidering the definition of IUP, their dates, and technological characteristics. In this paper, the IUP
assemblages are synthesized based upon currently published data illustrating their chronology anddistribution in northern China. By means of technological comparisons with assemblages in Siberian
Altai and Northern Mongolia, its regional significance of northern China IUP will be discussed as well.

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INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

New progress on the excavation and research of Shuidonggou site
complex________________________________________________________________________________Fei Peng1, Huimin Wang 2, Xing Gao3,4,5
1)

Department of Archaeology and Museology, School of Ethnology and Sociology, Minzu University

2)

Ningxia Institute of Archaeology and Cultural relics

3)

Laboratory for Vertebrate Evolution and Human Origins of CAS at the Institute of Vertebrate Paleontology andPaleoanthropology, Chinese Academy of Sciences

4)

CAS Center for Excellence in Life and Paleoenvironment

5)

University of Chinese Academy of Sciences________________________________________________________________________________
Shuidonggou site complex is one of most important Upper Paleolithic site in Northeast Asia. Since it
was discovered in 1923, a series excavation had been carried out. Totally 9 localities including SDG15, 7-9, 12 had been systematically excavated before 2007. Abundant material has demonstrated theculture diversity and evolutionary trajectory in Northwestern China before 10,000 to 50,000 years ago.
Meanwhile, a series issues about ancient human activities in this region still need more evidences from
SDG. Since 2014, new excavation on SDG1 and SDG2 which are the most important localities among
12 localities have been carried out. Numerous artifacts including pendant, ostrich eggshell beads,
blades, flakes etc and mammal fossils were yielded from several culture layers in these two localities.
Dozens of hearths were unearthed as well. Preliminary research has clarified the chronology of SDG2
and provide some important new information to investigate the onset of Upper Paleolithic in thisregion.

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Genetic insight into northern East Asia during the period of Ice Age andPaleolithic-Neolithic transition____________________________________________________________________________

Xiaowei Mao
Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences____________________________________________________________________________
Northern East Asia was inhabited by modern humans, marked by the Tianyuan man who was found at
the Tianyuan Cave near Beijing, as early as 40 ka. After 40ka, similar to Europe, northern East Asia
has been largely impacted during two important periods: the Last Glacial Maximum (LGM) and thetransition of Paleolithic to Neolithic (the beginning of agriculture). Using genome-wide data obtained
from individuals dated to 33.6-3.4 ka from the Amur Region, Liao River region, and Yellow Riverregion, we investigated what was the population history of northern East Asian and how did they adapt
to drastic changes of climate or lifestyle. For the population history, our investigation demonstrated
that Tianyuan and AR33K related ancestry was widespread in northern East Asia before the LGM. At
the end of the LGM stadial, the earliest coastal northern East Asian, who are basal to ancient northern
East Asians, appeared in the Amur Region. After 14 ka, ancient populations in the Amur Region
represent the closest East Asian source known for the Ancient Paleo-Siberians. After the PaleolithicNeolithic transition, we also observed more southern Asian related ancestry appeared in the ancient
populations in Yellow River region. For the adaptation, we observed that EDAR V370A mutation (anEast Asian specific adaptive variant which is associated with thicker hair shafts, more sweat glands,
and shovel-shaped incisors), was likely to have been elevated to high frequency after the LGM,
suggesting a possible timing for its selection. Our research provides the first look into the deep
population history and adaptation of northern East Asia.

44

INSIGHTS INTO HUMAN HISTORY IN THE EURASIAN STONE AGE

The Initial Upper Paleolithic in Asia: Is it still a useful concept?
________________________________________________________________________________

Steven L. Kuhn
School of Anthropology, University of Arizona________________________________________________________________________________The term Initial Upper Paleolithic (IUP) was originally applied to a single lithic assemblage in orderto describe its inferred position on a continuum from Middle to Upper Paleolithic.

The term waslater broadened to include a range of assemblages in the eastern Mediterranean, formerly called byother names, which shared certain elements of lithic technology. Subsequently the definition of the
IUP has been expanded even further to encompass assemblages distributed from the Jordan Valley to
the Siberian Altai, and from the Czech Republic to China.

While researchers have been very

successful in adding new archaeological assemblages to the corpus of the IUP, its sheer geographic
dispersal does beg the question of what kind of cultural phenomenon it could represent. Some scholars
consider the IUP to index the dispersal of anatomically modern Homo sapiens, whereas others see it
as simply the result of convergence in lithic technology at the end of the Middle Paleolithic. This
paper considers a number of alternative explanations in terms of current evidence. It also addresses
the following question: if the IUP phenomenon is not unambiguously an expression of shared culturalroots, is it still a useful thing to think about?

45

INTERNATIONAL SYMPOSIUM

Keynote Lecture

Pitfalls and progress in dating the earliest appearance of Homo sapiensin Central and Northeast Asia________________________________________________________________________________Tom Higham1,2
1) Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria2) Oxford Radiocarbon Accelerator Unit, University of Oxford, Oxford, UK________________________________________________________________________________
Chronology is crucial in understanding the movement and dispersal of Homo sapiens, and otherhominins, across Eurasia, as well as understanding their interactions and disappearance. Dating
material that is older than ~30,000 BP is challenging, but work undertaken over the last two decades
has improved the situation. My group has developed and improved several of the crucial pre-treatment
chemistry steps required to effectively decontaminate the most common archaeological materials fordating (bone, charcoal, shell), particularly the purification of bone collagen using ultrafiltration and in
the dating of single amino acids from collagen, as well quantifying background limits and corrections.
Within a large project funded by ERC, we have applied these methods to the analysis of over 1,000
samples of bone, shell and charcoal from more than 100 key Palaeolithic sites across Eurasia. The
main focus has been on sites with a succession of contexts containing lithic industries attributed to theMousterian (all mostly associated with Neanderthals) and the Initial and Early Upper Palaeolithic(thought to be related to Homo sapiens, but not always exclusively it would seem).
In this talk I will discuss the challenges in reliable radiocarbon dating, the importance of robust
pretreatment chemistry and Bayesian analysis using other complementary dating methods, and givesome initial results for the early dispersal of humans in this vast region of the world.

46

WORKSHOP 1

Emergence of regional diversityof Northeast Asia

WORKSHOP 1

Keynote Lecture

Regional differentiation from EUP to LUP in Northeast Asia:
An overview________________________________________________________________________________

Kelly E. Graf
Department of Anthropology, University of Kansas________________________________________________________________________________
Following the earliest appearance of Homo sapiens in Northeast Asia and after approximately 40 ka,
regional variation emerges quite quickly in the archaeological record with differences through timeand across space. From 40-25 ka we see decrease in the use of flat-face blade core technology with
development of various flake-based and macro-blade-based technologies, rise and fall of mobile art
production, change from opportunistic hunting by landscape novices to planned hunting andseasonally-based settlement by landscape experts, exploration of the extreme far north, and boating toprocure obsidian from insular extraction sites in the east. After 25 ka and during the LGM, sites acrossthe interior of this vast and varied region abate; however, along the eastern seaboard they continue andchange as wedge-shaped microblade core technologies develop and eventually spread inland to thewest and north. From 21-14 ka regional differentiation begins to take hold with continuation of only
microblade-based technologies in some areas, emergence of various bifacial technologies in others,
the combination of these two technologies in yet others, and development of a ceramic industry in the
Far East.
What led to regional differentiation and changes in technologies and land-use strategies after 40 ka?
Did this variability result from human responses to highly fluctuating climate and environmental
changes that affected distributions and availability of resources? Did they result from population
isolation and subsequent cultural identity development? These and similar questions will be exploredduring the lecture.

48

EMERGENCE OF REGIONAL DIVERSITY OF NORTHEAST ASIA

Upper Paleolithic exchange networks in Siberia and Mongolia________________________________________________________________________________

Arina M. Khatsenovich1, Evgeny P. Rybin1, Roman A. Shelepaev2, and John W.
Olsen1,3
1)Institute of Archaeology and Ethnography SB RAS, Novosibirsk, Russia

2)

V.S. Sobolev’ Institute of Geology and Mineralogy SB RAS, Novosibirsk, Russia

3)

School of Anthropology, University of Arizona, Tucson, USA________________________________________________________________________________
People began transporting exotic and/or high-quality raw material over long distances in Central Asia
beginning at least as early as the Final Middle Paleolithic. Rare cases of such transport have beenrecorded in the Russian and Gobi Altai Mountains, suggesting migrations and mobility rather thanexchange networks. The Initial Upper Paleolithic in Siberia and Mongolia was characterized by the
rapid spread of ideas, specific tool types, technologies, and non-utilitarian objects. Not all can be
explained as the result of human mobility. The most prominent examples include ornaments and nonutilitarian objects. While IUP ostrich eggshell bead-making in Mongolia and, probably, Transbaikalia,
was based on local raw material, the occupants of Denisova Cave in Siberia could acquire eggshell
only through exchange between populations or extremely long-distance transportation. Thisconclusion is supported by the sophisticated bone ornament-making apparent in Denisova Cave dueto the lack of softer and more easily worked ostrich eggshell. Highly variable stone raw material used
to fabricate ornaments in northern Mongolia reveals connections with the Russian Transbaikal Region.
The rapid spread of specific Initial Upper Paleolithic tool types throughout Central Asia and southern
Siberia is also associated with exchange networks and cannot be explained simply by migration.
Exotic raw material variability increased in EUP archaeological complexes. In some areas, such as
Transbaikalia and the Altai Mountains, this can be explained by improved knowledge of geologicalresources in populations’ home ranges, but also increased human mobility and the expansion of homeranges. Our research considers potential cases of long-distance contacts during the Initial and Early
Upper Paleolithic and a spectrum of fabrication methods, identifying some as evidence of exchange
networks rather than highly mobile human populations.

49

WORKSHOP 1

Paleolithic archaeology in the Qinling Mountains region, Central China________________________________________________________________________________Shejiang Wang
Institute of Vertebra Paleontology and Paleoanthropology, Chinese Academy of Science________________________________________________________________________________
Since 1995, we surveyed and excavated Paleolithic sites in the Qinling Mountains region, Central
China. The areas include the Luonan-Lushi Basin, Shangdan Basin, Hanzhong-Ankang Basin andLantian areas. The chronological data show that commencement of loess deposits in this region is atleast since two million years ago. The open-sites in the Luonan Basin which located at the second
terrace dated between 0.6-0.07 Ma. However, the sites which located on the second terraces in the
Hanzhong Basin, Lantian areas, and the sites located on the higher terraces in the Luonan Basin showsthat the loess deposits formed in the late Pleistocene. From the Early Pleistocene to the Middle
Pleistocene, it appears that the regional lithic assemblage belongs to the Oldowan (Mode I) lithic
industry, and it is dominated by choppers, cores, flakes, and simple retouched flake tools. However,from the Late Middle Pleistocene to the Late Pleistocene (between 250 ka to 50 ka), there were a large
number of Model II (Acheulean) lithic artifacts, such as hand-axes, picks, cleavers, spheroids, andknives found in this region, shows that the lithic industry had experienced an important transformation
process and the Acheulian LCTs lasted to the later Pleistocene in the region.

50

EMERGENCE OF REGIONAL DIVERSITY OF NORTHEAST ASIA

Keynote Lecture

The ebb and flow of human dispersals into Central Europe and further
North from the Last Glacial Maximum to the beginning of Late Glacial
warming, 25–14 cal kBP________________________________________________________________________________

Olaf Jöris 1,2,3
1)

Römisch-Germanisches Zentralmuseum – MONREPOS Archaeological Research Centre and Museum for
Human Behavioural Evolution, Neuwied, Germany

2)

Institute of Ancient Studies, Department of Prehistoric and Protohistoric Archaeology, Johannes Gutenberg
University, Mainz, German

3)

MOE Key Laboratory of Western China’s Environmental Systems Research School of Arid Environment andClimate Change, Lanzhou, China________________________________________________________________________________
Until the late 1990ies it was widely accepted that Central Europe had been void of humans during theLast Glacial Maximum (LGM), and that re-occupation took place no earlier than ~16–15 cal kBP, i.e.,during the Late Magdalenian. Over the last few decades, new radiocarbon dates and targeted dating
programs have changed this picture significantly, showing that Central Europe experienced repeatedphases of human presence and absense during the period ~25–14 cal kBP (Street & Terberger 2004;

Jöris & Street 2014; Kozłowski et al. 2017; Reade et al. 2020; Maier et al. 2020; Jöris 2021). At
least three discrete phases of human presence can be recognized in Central Europe over this timeinterval (Jöris & Street 2014): (1) sporadic occupation at ~23 cal kBP (Greenland Interstadial 2),
during which the southern half of Central Europe was occupied by populations that most likely arrived
from the East (Street & Terberger 2004; Reade et al. 2020); (2) a short episode of human incursion
into Central Europe at ~18.5 cal kBP, when mid-Magdalenian groups from France expanded eastwards,
eventually reaching southern Poland (Kozłowski et al. 2017); and (3) the major phase of Late
Magdalenian population expansion that started ~16 cal kBP. Present evidence indicates that Western
European populations expanded into Central Europe not simply by random dispersal, but steered bysystems of rules and regulations implemented by Late Magdalenian societies. Archaeological data
further indicate that this colonisation process spawned interactions between these Western European
populations and groups of Eastern European origin (Maier et al. 2020; Jöris 2021). The potentialsocial and technological exchanges between the two populations may have led to more sufficient
adaptations to overcome the challenges these Late Upper Palaeolithic hunter-gatherers faced in Central
Europe at a time of limited biomass production. This Late Magdalenian expansion represents thefoundation of a more northward migration at the transition to the Late Glacial interstadial (Greenland
Interstadial 1). It has been argued that the suceeding Central European Federmessergruppen (FMG),
i.e., curve-backed point industries, evolved from this Late Magdalenian substrate. However, it must
be discussed whether the FMG originated in the Late Epigravettian of Italy and south-eastern Europe.
51

WORKSHOP 1

A Late Epigravettian dispersal would represent a fourth phase (4) of northward population expansionthat crossed the Alps at ~14.3 cal kBP and accounts for a major turnover of the archaeological record
throughout the northern half of Europe. This interpretation is in line with a major population shift thathas been recognised for the Late Glacial period using aDNA (Bortolini et al. 2020). The earlier
dispersals and incursions into Central Europe may also be archived in past DNA signatures.
References
Bortolini, W., Pagani, L., Oxilia, G., Posth, C., Fontana, F., Badino, F., Saupe, T., Montinaro, F., Margaritora, D., Romandini, M.,
Lugli, F., Papini, A., Boggioni, M., Perrini, N., Oxilia, A., Aiese Cigliano, R., Barcelona, R., Visentin, D., Fasser, N., Arrighi,
S., Figus, C., Marciani, G., Silvestrini, S., Bernardini, F., Menghi Sartorio, J.S., Fiorenza, L., Moggi Cecchi, J., Tuniz, C.,
Kivisild, T., Gianfrancesco, F., Peresani, M., Scheib, C.L., Talamo, S., D’Esposito, M., Benazzi, S. 2020. Early Alpine
occupation backdates westward human migration in Late Glacial Europe. Current Biology 31, 1–10..
Jöris, O. 2021. The Magdalenian of Gönnersdorf and its Headless Anthropomorphic Depictions: On Social Coherence and the Late
Upper Palaeolithic Colonization of Central Europe. In: Gaudzinski-Windheuser, S., Jöris, O. (eds.) The Beef behind all Possible
Pasts. The Tandem-Festschrift in Honour of Elaine Turner and Martin Street. Monographien des Römisch-Germanischen
Zentralmuseums 157,1. Verlag des Römisch-Germanischen Zentralmuseums, Mainz, 325–368.
Jöris, O., Street, M. 2014. Eine Welt im Wandel. Die späteiszeitliche Besiedlungsgeschichte des Rheinlandes im Kontext von Klima
und Umwelt. In: LVR-Landesmuseum Bonn (ed.) Eiszeitjäger: Leben im Paradies? Europa vor 15000 Jahren. NünnerichAsmus, Mainz, 13–27.
Kozłowski, S.K., Terberger, T., Bobak, D., Orschiedt, J., PołtowiczBobak, M. 2017. Eastern borders of the Magdalenian ‘à navettes’
Maszycka cave in Lesser Poland (Southern Poland). In: Bourdier, C., Chehmana, L., Malgarini, R., Połtowicz-Bobak, M. (eds.),
L’essor du Magdalénien. Aspects culturels, symboliques et techniques des faciès à Navettes et à Lussac-Angles. Actes de laséance de la Société préhistorique française de Besançon, 17-19 octobre 2013. Société préhistorique française, Paris, 187–205.
Maier, A., Liebermann, C., Pfeifer, S.J. 2020. Beyond the Alps and Tatra Mountains – the 20-14 ka Repopulation of the Northern Midlatitudes as Inferred from Palimpsests Deciphered with Keys from Western and Central Europe. Journal of Paleolithic
Archaeology, 3, 398–452.
Reade, H., Tripp, J.A., Charlton, S., Grimm, S., Sayle, K.L., Fensome, A., Higham, T.F.G., Barnes, I., Stevens, R.E. 2020. Radiocarbon
chronology and environmental context of Last Glacial Maximum human occupation in Switzerland. Scientific Reports 10: 4694.
Street, M., Terberger, T. 2004. The radiocarbon chronology of the German Upper Palaeolithic: fifteen years of cooperation with ORAU.
In: Higham, T.F.G., Bronk-Ramsey, C., Owen, D.C. (eds.) Radiocarbon and Archaeology. Proceedings of the 4th Symposium,
Oxford 2002. Oxford University School of Archaeology Monograph 62. Oxford University School of Archaeology, Oxford,
281–302.

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EMERGENCE OF REGIONAL DIVERSITY OF NORTHEAST ASIA

Denisovans: Age, culture and habitat________________________________________________________________________________

Maxim Kozlikin
Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences________________________________________________________________________________
Denisova Cave is the oldest of the inhabited caves in Siberia and one of the most informative objects
for studying the ancient human culture and the natural environment around it, not only in North andCentral Asia, but also in the entire Eurasian continent.
Denisova Cave is located in the northwestern part of the Altai mountain system in the Anui river valley.
As an archaeological site the cave was discovered in 1977. The objects discovered testified that thecave had been inhabited since the Middle Paleolithic. From 1982 to the present, archaeologists fromthe Institute of Archeology and Ethnography of the SB RAS have been conducting research in the
Denisova Cave.
Denisova Cave became world famous in 2010, when the "Nature" journal published the results of agenetic analysis of human fossils found in the cave – a fragment of the terminal phalanx of the little
finger of a girl aged 6–7 years. Deciphering first mitochondrial and then nuclear DNA from this bone
sample showed that it belongs to a previously unknown hominin, which was named Denisovan fromthe place of discovery of anthropological remains.
As a result of interdisciplinary research, a unique stratigraphic sequence has been established, thanks
to which the continuous development of the cultural traditions of ancient human has been traced overthe past 300,000 years. Among the numerous Paleolithic materials from Denisova Cave, special
attention is drawn to the finds reflecting the spiritual and social aspects of the life of primitive man ornaments and objects of symbolic activity made of bone, mammoth tusk, animal teeth, ostrich egg
shells and soft stone. The main collection of these items was obtained from deposits 35–50 ka BP. It
includes a variety of pendants, beads, rings and bracelets. Currently, these are the most ancient
ornaments known in Eurasia.

53

WORKSHOP 1

10,000 years without Levallois: IUP industries of Ushbulak site, Eastern
Kazakhstan______________________________________________________________________
Anton Anoikin1, Kharevich Vladimir1, Pavlenok Galina1, and Taimagambetov
Zhaken21)

Institute of Archaeology and Ethnography SB RAS, Novosibirsk, Russia

2)

National Museum of the Republic of Kazakhstan, Nur-Sultan, Kazakhstan______________________________________________________________________
Ushbulak (ShiliktyValley, Eastern Kazakhstan) is a stratified site (8 layers): the Holocene assemblage(layer 1), the final UP assemblage (layers 2.1-3.3), the developed UP assemblage (layers 4-5.1), andthe initial UP assemblage (layers 5.2-7.2). Detailed age model for Ushbulak site was created using
Bayesian statistics on the basis of OSL -chronology and additional AMS ages. The assemblage fromthe lowermost layers 5.2-7.2 contains blade cores with the opposite platforms, numerous coretrimming elements, core-burins, end-scrapers on blades, including those with ventral base thinning,
truncated-faceted tools, truncated blades, a biface and an oblique point. Core trimming elements
correspond well to the available cores. Experimental and attributive approaches established theabsolute predominance of subprismatic bidirectional laminar method, based on well-prepared precores knapping. Based on the composition of the lithic industry, layers 5.2-7.2 can be defined as alithic workshop at the outcrops of raw material. The Ushbulak lithic industry is similar to the stratified
assemblages attributed to the IUP in Southern Siberia (Kara-Bom site and others) and Northern
Mongolia (Tolbor-4 site and others).
The most pronounced traits that make Ushbulak distinctive to IUP industries of neighboring regions,
are total dominance of the only method for bidirectional blade production and total lack of Levallois
component in the technocomplexes. Along with that, any changes in primary knapping methods andtool assemblages have not been revealed in IUP complexes of Ushbulak, nonetheless its long-termpresence in the region (~10 ka). This industry first appeared in Eastern Kazakhstan about 49–47 ka in
its complete form and existed changeless up to its disappearance about 37–35 ka.
Acknowledgements
This study was supported by the Russian Science Foundation (21-78-10146).

54

WORKSHOP 2

Recovering ancient remains
and reconstructing past

WORKSHOP 2

Function and behaviour: Recognizing use-wear on prehistoric tools________________________________________________________________________________

Simona Arrighi1,21) Department of Cultural Heritage, University of Bologna2) Department of Environment, Earth and Physical Sciences, University of Siena________________________________________________________________________________
Stone tools are a primary source of information to reconstruct behavioural evolution of hominin
species, including subsistence activities, social organization, and symbolic thinking. Therefore,
understanding the function of prehistoric tools is a pivotal marker of ancient behaviours and activities.
Use-wear analysis is a methodology used to identify the function of stone tools as well as the functionof tools made on other materials such as bone, ivory, etc… This discipline, which has been introduced
for the first time in a codified way by S.A. Semenov in the 1930s, is based on the observation ofphysical alterations produced by use on the working edges and/or surfaces of artefacts. Suchalterations are usually interpreted by means of comparison with traces experimentally produced
reproducing different activities with tool replicas.
Use-wear analysis is carried out using different types of microscopes and a range of magnifications inrelation to different techniques of observation: low-power approach (LPA) and high-power approach(HPA), commonly used in combination.
LPA is conducted by way of a stereomicroscope with magnifications ranging from 10x to 50x. Thismethod of observation allows us to record edge rounding and edge removals as wear traces that makeit possible to identify the hardness of the processed material and type of action carried out (e.g., cutting,scraping…). HPA is carried out using a reflected light microscope (with 50x-500x magnifications) andit allows us to recognize microwear such as polished areas and striations along with micro-roundingof the edge. By means of HPA it is possible not only to distinguish the degree of hardness of theworked material, but also to identify different types of materials (e.g., hide, wood, bone, etc). In thelast decades many studies focused on developing techniques that use quantitative methods to bettercharacterize use-wear, such as, for example, laser scanning confocal microscope.

56

RECOVERING ANCIENT REMAINS AND RECONSTRUCTING PAST

Methodology for the use-wear analysis developed by Tohoku University
Microwear Research Team (TUMRT)________________________________________________________________________________

Yoshitaka Kanomata
Graduate School of Arts and Letters, Tohoku University________________________________________________________________________________
Tohoku University Microwear Research Team was originally organized by Professor Chosuke
Serizawa, one of the pioneers in the field of Japanese Palaeolithic studies. In 1976, he started to apply
the high-power microwear analysis, and organized the research team including his students. The firstseries of experiments indicated that precise interpretation should be done by combining several
categories of use-wear data such as polish, striation and microflaking. Therefore, in order to evaluate
use-wear traces comprehensively, both “high-power” and “low-power” methods should be applied.
By 1982, his team developed micro-polish classification, based on experimental program with
abundant siliceous hard shale, chert, and obsidian. They classified features of the polished surface
based on characteristics attributes such as brightness, smoothness, roundness, extension, elevation,connection, pit-features, and-so-on. This classification became an index for lithic functional analysis
in Japanese archaeology. At the moment, 11 types of polish are used as criteria for the interpretationof use-wear.
As another pioneering study, Professor Kaoru Akoshima introduced the concept of the organization oftechnology and the middle-range theory (Binford 1979) to Japanese archaeology: these concepts wereapplied to the experimental microwear studies as a theoretical framework.
Practical studies of microwear analyses were carried out for the lithic artifacts in the Upper Palaeolithic,
Jomon and Yayoi periods, in Japan. These studies led to understand the relationships between typology
and functional aspects of lithic tools. The functional variability, re-sharpening, reuse, and degree ofcuration depend on the technological organization of prehistoric hunter-gatherers and farmers.
This presentation focuses on the methodological problems and research achievements TUMRT
through the 45-year of research.

57

WORKSHOP 2

Methods for identifying hunting traces and its application for earlyUpper Palaeolithic sites in Japan________________________________________________________________________________

Shunsuke Totsuka1 and Katsuhiro Sano2
1) Gradual School of Arts and Letters, Tohoku University
2) Center for Northeast Asian Studies, Tohoku University________________________________________________________________________________
A stone tip attached onto a wooden spear produces a variety of fractures when they hit an animal prey.
While some impact fracture types can occur due to other factors, including blank flaking, retouching,
and trampling, there are also specific fracture patterns that are yielded exclusively from hunting. Oneof the important clues for identifying impact fractures is bending initiation. Different from intentional
flaking on lithics that produces a cone scar, impact fractures occur by bending force throughlongitudinal collision with an animal target and exhibit no negative bulb on the surface. However,spin-off fractures, which are secondary fractures caused by contact between the two primarily formed
broken surfaces of a stone tip, often show a cone at the initiation (but they are sometimes bending
fractures). Therefore, while primary impact fractures exclusively initiate with bending, spin-off
fractures frequently show a negative bulb on the surface. In addition to these principal characteristics,
our experiments and those of others have demonstrated that flute-like, burin-like, bifacial spin-off, andsingle spin-off fractures longer than 6 mm are reliable as diagnostic impact fractures (DIFs). These
fracture types seldom occur due to the other factors. Transverse fractures with step, hinge, and feather
terminations can be regarded as DIFs only if these fractures form after lateral retouching. Based on
the reliable DIFs, we examined early Upper Palaeolithic (EUP) assemblages in the Kanto region of
the Japanese islands to identify hunting armatures. The results of a systematic microfracture analysisof the EUP assemblages are presented and future issues to evaluate the ratio of DIF occurrences among
different raw materials are discussed.

58

RECOVERING ANCIENT REMAINS AND RECONSTRUCTING PAST

Quantifying morphological variation: Applications of geometricmorphometrics to archaeological artifacts________________________________________________________________________________

Kohei Tamura1, 2
1)

Frontier Research Institute for Interdisciplinary Sciences, Tohoku University

2)

Center for Northeast Asian Studies, Tohoku University________________________________________________________________________________
Analyzing morphological variation is an important step to extract information from archaeologicalremains. Computational methods have been increasingly used to quantify morphological variation in
an archaeological data set, at least partly because of the increased access to large data sets of digitized
three-dimensional data and the trends of reproducible research and open science. In addition, somescholars have emphasized, as an advantage of such methods, the importance to calculate relative‘similarity’ values among individual artifacts rather than dividing the focal set of artifacts to somecategories and assuming morphological homogeneity in each category.
Geometric morphometrics, originally developed in evolutionary biology and biological anthropology,
is a suite of such quantitative methods and have been increasingly used to quantify morphologicalvariation in archaeology. Examples range from stone tools to monuments. Advantages of geometricmorphometrics approaches over ‘traditional’ ones include that geometric information is modelled as
relative positions of coordinates, and thus there are a wide variety of ways to visualize morphologicalvariation.
In this talk, I will first illustrate the concept of geometric morphometrics approach and demonstratesome simple examples in archaeology using R, including both outline- and landmark-based methods.
In addition, I will briefly introduce some advanced applications in archaeological research including
examples using digitized three-dimensional models. The limitations of geometric morphometrics inarchaeological research will be also discussed.

59

WORKSHOP 2

Approaching Upper Palaeolithic hunting strategies through technology
and residue analysis________________________________________________________________________________

Matteo Rossini1 and Clarissa Dominici1
1)

Department of Environment, Earth and Physical Sciences, University of Siena________________________________________________________________________________
Backed tools are one of the most abundant categories of retouched blanks in Upper Palaeolithic
assemblages and play a key role in the definition of specific cultural techno-complexes, possibly
showing great variability in time and space. For this reason, their study is fundamental to understandbehavioural dynamics of Palaeolithic hunter-gatherers. Based on archaeological and ethnographic
cases, as well as on the numerous wear studies that have been produced in recent years on these
artefacts, it can now be assumed that they were mainly used as part of composite weapon systems.
The study of backed tools through a modern integrated approach combining technological,
morphometric and residue analysis enables us to greatly increase our understanding of thetechnological know-how related to hunting strategies. This kind of approach allows us to answer threemain questions: 1) how armatures were manufactured, 2) how they were used and, eventually, 3) therole (cultural, environmental, chronological etc.) these specific artefacts have in defining the technoeconomic behaviour of different groups.
For what concerns manufacturing modalities, an important role is played by the kind of backing
technique used. It represents a fundamental aspect of the transformation of blanks in backed tools andhas recently been investigated in experimental and methodological works, as well as archaeologicalcase studies.
Use, on the other hand, can be inferred through residue analysis, which can provide important
information on the types of adhesives selected to fix lithic implements to the shaft, as well as on theirrelative position when combined with functional analysis of diagnostic impact fractures. Residue
analysis also deals with the identification of organic and inorganic substances with which lithic
implements have come into contact during use, namely animal tissues, vegetal fibres etc., thus
representing an essential discipline in the study of hunting strategies.

60

RECOVERING ANCIENT REMAINS AND RECONSTRUCTING PAST

Zooarchaeology and the Middle to Upper Palaeolithic transition:
Some methodological insights________________________________________________________________________________

Francesco Boschin1 and Jacopo Crezzini1
1)

Department of Environment, Earth and Physical Sciences, University of Siena________________________________________________________________________________
Animal remains from Palaeolithic contexts offer the possibility of exploring human-environment
interactions and subsistence economy of hunter-gatherer groups. An intriguing topic is the so called“Middle to Upper Palaeolithic Transition” in Europe, a period in which not only a cultural turnover
took place, but also the replacement of human populations (Neanderthals) due to the arrival of newgroups with their own biological characteristics and needs (Modern Humans). Zooarchaeology canhelp us to disentangle changes in strategies for exploiting animal resources, not only considering preyselection in terms of species, but also in terms of age classes and anatomical parts. Despite thepotentialities that this field of research offers, Palaeolithic zooarchaeology has to face peculiar
problems, such as a high level of bone fragmentation, low taxonomic and anatomical identifiability of
specimens and the presence of several actors who can alter (or create) the bone samples (e.g., hyaenas).
Peninsular Italy offers the possibility to study in depth the demise of Neanderthals and the rise ofModern Humans from a zooarchaeological perspective. The talk will focus on case studies from cavesites, highlighting the methodological issues we must consider in order to obtain the best results is thisparticular field of research. Attention will be paid to the environmental influence on game availability,
the identification of bone accumulators (distinguishing human camps from hyaena dens or mixed
samples) and the influence of hunting strategies on prey selection. In addition, the different
significance of peculiar animal resources for different human groups will be explored.

61

WORKSHOP 2

Disentangling the spatial structure of a Palaeolithic camp:
Methodological issues________________________________________________________________________________

Vincenzo Spagnolo
Department of Environment, Earth and Physical Sciences, University of Siena________________________________________________________________________________The spatial structure of a Palaeolithic site is a wide window on multiple information about thesettlement strategies of hunter-gatherer groups. Nevertheless, assuming the existence of a direct
correspondence between social, behavioural and spatial structures at an archaeological site is clearlynaive. For this purpose, both a high level of interdisciplinary and a complex and highly specialized
perspective (the spatial archaeologist’s one) are required. Nowadays the Spatial Archaeologists canbenefit from very powerful investigative tools: the GIS. It acted a real technological andepistemological revolution, but the actual implications of this scientific debate are not yet expressedto their full potential. The GIS is not simply a software aimed at creating maps, it is the most developed
expression of both philosophical and technical merging of spatial sciences. In particular, it has madethe field of Spatial Archaeology the natural terrain expressing the best interdisciplinary convergenceof contextual and behavioural approaches. A robust analytical protocol is required to best express thisnature, correctly responding to the epistemological issues triggered by modern Spatial Archaeology.
Here, we present the work-flow developed for the intra-site study of palaeolithic sites, it includes 4
steps: 1) definition data and geodatabase design, 2) taphonomic analysis of the context, 3) spatialfunctional analysis, 4) reconstruction of the activity areas and functional structure of the site. The firststep, in particular, plays the decisive role, because the goodness of data definition and classification
directly influences the “quality” the results.

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RECOVERING ANCIENT REMAINS AND RECONSTRUCTING PAST

Lithic technology, the notion of chaîne opératoire and its application in
two case studies: The Mousterian of Riparo l’Oscurusciuto and theUluzzian of Grotta di Castelcivita_____________________________________________________________________________

Giulia Marciani 1,2
1)

Department of Cultural Heritage, University of Bologna

2)

Department of Environment, Earth and Physical Sciences, University of Siena_____________________________________________________________________________The notion of chaîne opératoire – first conceived by Leroi-Gourhan – is a method used to retrace thereduction sequence used in the production of stone artefacts. This entails arranging the actions required
to envision and create the lithic tools in spatiotemporal order. Thus, the stone tool production process
can be divided into several phases: the selection and acquisition of the appropriate raw material; theinitialization and structuration of the raw block of material in order to produce stone tools withpredetermined shape and technical features; the production of target flakes; the further maintenanceof the convexities in order to produce other desired target objects; the eventual transformation by
retouch of the products; and finally, the use of the tools. The tools could then be either abandoned orreused and recycled at the end of this process. Reconstructing the reduction sequence of anarchaeological assemblage allows us to learn about the prehistoric craftsmen's intentions, the gesturesand procedures involved in manufacturing the tools, and how they were used. We can thereby rebuilda series of complex behaviours from a single object.
The purpose of this talk is to illustrate the lithic reduction sequences of two Southern Italian sites: theOscurusciuto rockshelter and Grotta di Castelcivita. Oscurusciuto is an important Mousterian site, andit contains a long succession of levels dating from the end of the middle Palaeolithic (42,724 ± 716
cal BP and 55 ± 2 ka 40Ar/39Ar). The reduction sequence at the site is characterized by a dominating
Levallois production aimed at producing elongated and convergent tools. The second case study is
Castelcivita, which has an extended stratigraphy that covers the Mousterian, Uluzzian, andProtoaurignacian techno-complexes. The Uluzzian documents one of the earliest Homo sapiens
dispersions in southern Europe. In terms of technology, it represents a notable shift from the preceding
and broadly contemporaneous Mousterian techno-complexes. At Castelcivita, the Uluzzian
assemblages were thoroughly investigated and understood from a technological and functional
standpoint.

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WORKSHOP 2

Elemental and isotopic analyses of tooth enamel disclose individuals’ life
histories________________________________________________________________________________

Federico Lugli 1,2
1)

Department of Cultural Heritage, University of Bologna

2)

Department of Chemical and Geological Sciences, University of Modena________________________________________________________________________________
Recent advancements in mass spectrometry techniques allowed to gather more and more data on
animals’ and humans’ life history. This latter is broadly defined as the individual’s pattern of allocationof time and energy to various fundamental tasks (e.g. growth, reproduction, and survival). Specifically,
analyzing the elemental and isotopic composition of human and animal skeletal remains it is possible
to infer about their mobility pattern, their eating habits, their health status and their metabolism. Beinga growing tissue resistant to diagenetic alteration, tooth enamel is the foremost target for chemical
analyses, preserving a wealth of biogenic information on past life events for millions of years. Overall
this body of information may provide precious insights into evolutionary trajectories and adaptationof extinct and extant taxa. For more recent periods, individuals’ life histories can inform on societal
structures and cultural practices of people. With this lecture, I will give an overview of the current
elemental and isotopic methodologies applied to skeletal remains, presenting e.g. the application of(non-)traditional isotope systematics to trophic niche reconstruction; the measure of time-resolved
chemical data to infer about early dietary changes, mobility and exposure to toxic metals; the use ofdata modelling to formalize and maximize the collected information.

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Author Index
Adriana Moroni .................................................................................................................................... iv, vii, 25
Andrea Columbu ....................................................................................................................................... iii, 19
Andrea Picin .............................................................................................................................................. iv, 23
Anton Anoikin ............................................................................................................................................vi, 54
Arina Khatsenovich ........................................................................................................................... v, vi, 41,49
Bo Li ............................................................................................................................................................ iii, 9Chuan-Chou Shen....................................................................................................................................... iii, 9Clarissa Dominici ..................................................................................................................................... vii, 60
Cosimo Posth ............................................................................................................................................ iii, 20
Dongju Zhang ............................................................................................................................................. iii, 9Evgeny Rybin .................................................................................................................................... v, vi, 41,49
Fahu Chen ................................................................................................................................................... iii, 9Federico Lugli .......................................................................................................................................... vii, 64
Fei Peng....................................................................................................................................................... v, 43Feng Li ........................................................................................................................................................ v, 42Francesco Boschin.................................................................................................................................... vii, 61
Frido Welker .......................................................................................................................................... iii, 9, 13
Fumie Iizuka............................................................................................................................................... iv, 33
Giulia Marciani ........................................................................................................................................ vii, 63
Hiroki Oota ................................................................................................................................................ iv, 34
Huimin Wang .............................................................................................................................................. v, 43
Israel Hershkovitz ..................................................................................................................................... iii, 17
Jacopo Crezzini ........................................................................................................................................ vii, 61
Jean-Jacques Hublin.............................................................................................................................. iii, 9, 16
Katerina Douka ......................................................................................................................................... iii, 11
Katsuhiro Sano .......................................................................................................... i, ii, iv, v, vi, vii, 25, 31, 58
Kazuki Morisaki ................................................................................................................................... iv, 31, 32
Kelly Graf ...................................................................................................................................................vi, 48
Kharevich Vladimir ....................................................................................................................................vi, 54
Kohei Tamura................................................................................................................................... ii, iv, vii, 59
Kseniya A. Kolobova ........................................................................................................................... ii, v, 2, 41
Lisa Vaccari ................................................................................................................................................ iv, 25Masami Izuho ..................................................................................................................... ii, iii, iv, v, vi, 31, 39

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Matteo Rossini ......................................................................................................................................... vii, 60
Maxim Kozlikin ..........................................................................................................................................vi, 53
Nicolas Zwyns ............................................................................................................................................. v, 40
Olaf Jöris ............................................................................................................................................ii, vi, 5, 51
Marcel Weiss ................................................................................................................................................ ii, 5Olsen J.W ...................................................................................................................................................vi, 49
Pavlenok Galina .........................................................................................................................................vi, 54
Qiaomei Fu ................................................................................................................................................. iii, 9Sahra Talamo.............................................................................................................................................. iv, 26
Seiji Kadowaki ........................................................................................................................................... iii, 18
Shejiang Wang............................................................................................................................................vi, 50
Shelepaev R.A ............................................................................................................................................vi, 49
Shunsuke Totsuka .......................................................................................................................... iv, vii, 31, 58
Simona Arrighi ............................................................................................................................... iv, vii, 25, 56
Stefano Benazzi .......................................................................................................................... ii, iv, vii, 24, 25
Steven Kuhn ................................................................................................................................................ v, 45
Svante Pääbo ............................................................................................................................................... iii, 9Taimagambetov Zhaken .............................................................................................................................vi, 54
Ted Goebel .................................................................................................................................................. v, 38
Tom Higham ............................................................................................................................................... v, 46
Tsenka Tsanova ......................................................................................................................................... iii, 21
Vincenzo Spagnolo................................................................................................................................... vii, 62
Viviane Slon ............................................................................................................................................. ii, 3, 4
Xiaowei Mao................................................................................................................................................ v, 44
Xing Gao .......................................................................................................................................... iii, v, 10, 43
Xing Song ..................................................................................................................................................... ii, 8
Yoshihiro Nishiaki ........................................................................................................................................ ii, 7
Yoshitaka Kanomata ................................................................................................................................. vii, 57
Yousuke Kaifu ............................................................................................................................................ iv, 30

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