RNMH2014
The Second International Conference November 30 – December 6, 2014, Date City
Edited by Takeru Akazawa & Yoshihiro Nishiaki

RNMH PROJECT SERIES No. 003

RNMH2014
The Second International Conference

Replacement of Neanderthals by Modern Humans:
Testing Evolutionary Models of Learning

November 30 December 6, 2014, Date City

Program and Abstracts
Edited by

Takeru Akazawa &Yoshihiro Nishiaki

RNMH2014
The Second International Conference

Replacement of Neanderthals by Modern Humans:
Testing Evolutionary Models of Learning

November 30 December 6, 2014, Date City

Program and Abstracts
Edited by

Takeru Akazawa &Yoshihiro Nishiaki

RNMH Project Series
Editor-in-Chief

Takeru Akazawa (Kochi University of Technology)
Editorial Board

Shunichi Amari (Brain Science Institute)
Kenichi Aoki (The University of Tokyo)
Ofer Bar-Yosef (Harvard University)
Ralph L. Holloway (Columbia University)
Shiro Ishii (Japan Society for the Promotion of Science)
Tasuku Kimura (The University of Tokyo)
Yoshihiro Nishiaki (The University of Tokyo)
Naomichi Ogihara (Keio University)
Hiroki C. Tanabe (National Institute for Physiological Sciences)
Hideaki Terashima (Kobe Gakuin University)
Minoru Yoneda (The University of Tokyo)

Published by

RNMH Project Group

Research Institute, Kochi University of Technology

CIC Tokyo, 3-3-6 Shibaura, Minato-ku, Tokyo
108-0023, Japan
Tel: +81-(0)3-5440-9039
Fax: +81-(0)3-5440-9119
http://www.koutaigeki.org/

The publication of the RNMH Project Series has been
supported by a Grant-in-Aid for Scientific Researchon Innovative Areas (Grant No. 22101001) fromJapanese Ministry of Education, Culture, Sports,
Science and Technology.
ISBN: 978-4-9906371-0-1© 2014 RNMH Project GroupAll rights reserved. No parts of this publication may be
reproduced, stored in a retrieval system, or transmitted,
in any form or by any means, electronic, mechanical,
photocopying, recording, or otherwise, without the priorpermission of the RNMH project group.

Produced by Akita Kappan, Inc., Akita

CONTENTS
006 Preface
009 Conference Program
Abstracts – Oral Presentations

017 DAY 1 Opening Remarks
Keynote Lectures

025 DAY 2 Neanderthals and Modern Humans: Behavioral
Characterization
The First RNMH Processes in West Asia

055 DAY 3 Environmental Backgrounds for RNMHTiming of the RNMH in Europe
Global Processes of RNMH

DAY 4 Excursion (optional)

079 DAY 5 Learning Strategies and Cultural Evolution
105 DAY 6 Cognition and Learning

Looking into the Neanderthal Brain
Closing Remarks

DAY 7 Post-conference Excursion (optional)

127 Abstracts – Poster Presentations
179 Author Index

Preface
RNMH2014 is the second international conference for our major research project entitled “Replacement of
Neanderthals by Modern Humans: Testing Evolutionary Models of Learning” (RNMH). Similar to the first
conference held in Tokyo, Japan in 2012, this one aims to compile the latest research investigating the issues
surrounding the replacement of those two populations.
Understanding this phenomenon of replacement, including clarification of its detailed spatio-temporal processes
and interpretation of the mechanism(s) behind them, has been one of the most hotly debated subjects in the
recent anthropological literature. Even in the few years directly after launching the RNMH project in 2010, a
series of influential findings were newly reported. For example, particularly remarkable achievements were basedon ancient genetic studies. DNA analyses of fossil and Modern Human remains have demonstrated the possible
occurrence of interbreeding between Neanderthals and anatomically Modern Humans in the Paleolithic period.1
Likewise, the presence of another indigenous population in Paleolithic Eurasia, the Denisovans, was identified ineastern Central Asia where the occurrence of Neanderthals was also known.2 On the other hand, archaeologicalfieldwork has also continuously yielded new insights. Discoveries indicating “modern” Neanderthal behaviors thatwere previously attributed only to anatomically Modern Humans are at most interesting,3 blurring the behavioral
distinction between these two populations.
These and other new data indicate that the replacement processes should be considered more complicated than
before, certainly rejecting a simple straightforward “replacement” model. At the same time, varied processes,
and hence varied mechanisms in each region should be considered. In other words, research in this area today
requires more refined perspectives grounded in interpretation of higher resolution data. This recognition is related
to one of the two main objectives of this conference, which is giving an opportunity for international colleagues
to gather and present state-of-the-art results on these subjects. This opportunity should also be welcomed by
RNMH project members to present their research results in an international arena to solicit expert evaluation.
The second objective of this conference is to discuss the working hypothesis proposed by the RNMH project,coined as the “learning hypothesis.” The specific goal of this project is to verify this hypothesis within aninterdisciplinary research framework incorporating new perspectives and methods from the humanities andbiological sciences. It is unique in focusing on possible differences in learning (abilities/strategies/behaviors)
between Neanderthal and Modern Human societies. The hypothesis was based on the following reasoning:
success or failure of any human society is heavily dependent on culture; culture is transmitted, maintained, andmodified through learning; and this premise is probably applicable to human societies during the replacement
period as well. Was there any significant difference in learning between Neanderthals and anatomically Modern
Humans? If so, clarification is required to explain how and whether it derived from their innate biologicaldifference, social environments, or from both factors. Although the detailed answers will be obtained after muchresearch to come, we believe that it is a significant advantage to look at the broader replacement processes, thus
enlarging the dimensions of current debates.

006

Spanning seven days between November 30 and December 6, the conference consists of three major parts.
After the opening remarks and keynote lectures on Day 1 (November 30), when the latest findings in physicalanthropology, cognitive archaeology, and field archaeology are addressed, the first part starts on Day 2 (December
1) and continues until Day 3 (December 2). It deals with the behavioral characterization of these two populations
and the details of replacement processes by region as revealed through a variety of disciplines including
prehistoric archaeology, ecology, genetics, and palaeoenvironments. The regional survey focuses on West Asia andEurope, where the processes have been most intensively studied, but it also refers to new data from Africa andCentral and East Asia. Following a break for the excursion on Day 4 (December 3), the second subject, learning,
will be covered on Day 5 (December 4). Possible differences in learning abilities, strategies, and behaviors
between Neanderthals and Modern Humans are discussed using evidence from fieldwork in archaeology, cultural
anthropology, and primatology. Theoretical modeling to predict evolutionary paths in learning strategies will also
be addressed. In the final part on Day 6 (December 5), sessions will be devoted to discussing the innate learning
capabilities among these hominin populations. Instead of considering behavioral evidence reflected in thearchaeological records, the discussion focuses on independent evidence obtained through innovative approaches
to studying the available cranial endocasts of Neanderthals and anatomically Modern Humans. The conference
closes after the post-conference excursion on Day 7 (December 6).
RNMH2012, Tokyo, consisted of 30 oral and 38 poster presentations. RNMH2014 incorporates 55 oral and22 poster presentations. In addition to changes in the conference structure and number of presentations, we arepleased to see the remarkable increase in the range of countries in which participants are based (6 to 16). Thiscertainly reflects the worldwide increase in research interest on these topics. The organizers are deeply indebted to
all participants, especially those visiting from overseas who made RNMH2014 a truly international platform.
Last but not least, we would like to mention that financial support for RNMH2014 has been obtained from the
Ministry of Education, Culture, Sports, Science and Technology, Japan (Grant No. 22101001) and the Education
Board of Date City, Hokkaido. Our deepest gratitude is extended to the Date Volunteer Society for Scientific
Meeting, who organized the venue for this international meeting.
Takeru Akazawa

Kochi University of Technology

Yoshihiro Nishiaki
The University of Tokyo

1. Green RE et al (2010) A draft sequence of the Neandertal genome. Science 328: 710–722.
2. Krause J et al (2010) The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. Nature 464: 894–897. Reich D et al (2010)
Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature 468: 1053–1060.
3. Villa P, Roebroeks W (2014) Neandertal demise: An archaeological analysis of the Modern Human superiority complex. PLoS One. doi: 10.1371/journal.
pone.0096424

007

RNMH2014

The Second International Conference

Replacement of Neanderthals by Modern Humans: Testing Evolutionary Models of Learning
ORGANIZING COMMITTEE
Chair, Organizing Committee

Takeru Akazawa, Kochi University of Technology, Japan

Chair, Program Committee

Yoshihiro Nishiaki, The University of Tokyo, Japan

Committee Members

Shunichi Amari, RIKEN Brain Science Institute, Japan
Kenichi Aoki, Meiji University, Japan
Tomoya Aono, Date City Institute of Funkawan Culture, Hokkaido, Japan
Ofer Bar-Yosef, Harvard University, USA
Ralph L. Holloway, Columbia University, USA
Shiro Ishii, Japan Society for the Promotion of Science, Japan
Tasuku Kimura, The University of Tokyo, Japan
Naomichi Ogihara, Keio University, Japan
Naoyuki Ohshima, Date City Institute of Funkawan Culture, Hokkaido, Japan
Hiroki C. Tanabe, Nagoya University, Japan
Hideaki Terashima, Kobe Gakuin University, Japan
Minoru Yoneda, The University of Tokyo, Japan

SECRETARIAT
Yoshihiro Nishiaki, The University of Tokyo, Japan
Mayumi Yaguchi, Kochi University of Technology, Japan
Yayoi Ogawa, The University of Tokyo, Japan
Noriko Kagaya, Quality Management, Japan
Seiji Kadowaki, Nagoya University, Japan
Katsuhiro Sano, The University of Tokyo, Japan
Yasuhisa Kondo, Research Institute for Humanity and Nature, Japan
Chie Akashi, The University of Tokyo, JapanHEAD OFFICE RNMH

CIC Tokyo, 3-3-6 Shibaura, Minato-ku, Tokyo 108-0023, Japan
Email: humis@nii.ac.jp
Tel: +81-(0)3-5440-9039
Fax:+81-(0)3-5440-9119
URL: http://www.koutaigeki.org

008

CONFERENCE PROGRAM
DAY 1: November 30 (Sunday), 2014
11:00-17:00

Registration

Opening Remarks
13:00-13:40
Takeru Akazawa ……………………………………………………………………………………………………018
Will we Modern Humans fade away or be replaced?
Keynote Lectures

Chair: Yoshihiro Nishiaki

16:25-16:30
16:30-17:00
17:30

Silvana Condemi …………………………………………………………………………………………………019
From Neandertals to Modern Humans: A view from the Italic PeninsulaThomas Wynn and Frederick L. Coolidge ……………………………………………………………………021
The expert performance model of Neandertal cognition
Coffee breakMarcel Otte…………………………………………………………………………………………………………023
Neandertal: Abilities and realisations
Break
Discussion
Free bus service to the Toyako Manseikaku Hotel Lakeside Terrace

Welcome party
18:00-19:30

Toyako Manseikaku Hotel Lakeside Terrace

13:40-14:30
14:30-15:20
15:20-15:35
15:35-16:25

DAY 2: December 1 (Monday), 2014
9:30-17:00

Registration

Neanderthals and Modern Humans: Behavioral Characterization
Chair: Seiji Kadowaki

10:00-10:25

10:25-10:50
10:50-11:15
11:15-11:40
11:40-12:05

12:05-13:50

Michael Chazan, Francesco Berna, Steven Walker, Ian Watts, Jayne Wilkins andLiora Kolska Horwitz ……………………………………………………………………………………………026
The development of technology during the Earlier Stone Age: Research at Wonderwerk Cave andthe Kathu Complex, Northern Cape Province, South Africa
Ran Barkai …………………………………………………………………………………………………………029
A tale of three caves: The role of mammoths in Neanderthal adaptation and lifeways
Avraham Ronen ……………………………………………………………………………………………………031
Middle Palaeolithic flint mines in Mount Carmel: An alternative interpretation
Gonen Sharon ……………………………………………………………………………………………………034
A week (?) in the life of the Mousterian hunter
Taro Yamauchi ……………………………………………………………………………………………………036
Evolution of learning capacity and acquired behaviors of Modern Humans from the perspective oflife history and brain development
Lunch/ Tour to the Historical Museum of Date (optional)

009

The First RNMH Processes in West Asia
Chair: Katsuhiro Sano

13:50-14:15

14:15-14:40

14:40-15:05
15:05-15:30
15:30-15:50

Mina Weinstein-Evron, Yossi Zaidner, Alexander Tsatskin, Reuven Yeshurun and
Israel Hershkovitz …………………………………………………………………………………………………038
The Early Middle Paleolithic of Misliya Cave, Mount Carmel, Israel
Yossi Zaidner, Laura Centi, Marion Prevost, Maayan Shemer and Oz Varoner …………………………039
Preliminary study of the lithic assemblages from Nesher Ramla: A case of continuity andchange in the Levantine Middle Paleolithic
Andrew W. Kandel and Ron Shimelmitz ………………………………………………………………………040
Technological trends of the Middle to Upper Paleolithic transition at Sefunim Cave, Israel
Nigel Goring-Morris and Anna Belfer-Cohen …………………………………………………………………043
The Ahmarian in the context of the earlier Upper Palaeolithic in the Near East
Coffee break

Chair: Yasuhisa Kondo

15:50-16:15
16:15-16:40
16:40-17:05

17:05-17:30

18:00

Ofer Marder, Israel Hershkovitz and Omry Barzilai …………………………………………………………044
The Upper Palaeolithic of Manot Cave, western Galilee, Israel
Andreas Taller, Boris Gasparyan and Andrew W. Kandel ……………………………………………………045
Living on the edge: The earliest Modern Human settlement of the Armenian Highlands
Fereidoun Biglari …………………………………………………………………………………………………048
Typo-technological analysis of the Late Middle Paleolithic bifacial industry of
Qaleh Bozi Rockshelter, Central Iran
Sonia Shidrang ……………………………………………………………………………………………………051
The Middle to Upper Paleolithic transition in Zagros: The appearance and evolution of Baradostian in
the light of new research
Free bus service to the Toyako Manseikaku Hotel Lakeside Terrace

DAY 3: December 2 (Tuesday), 2014
9:30-17:00

Registration

Environmental Backgrounds for RNMH
Chair: Minoru Yoneda

10:00-10:25
10:25-10:50
10:50-11:15
11:15-11:40

010

Wonsuh Song, Yasuhisa Kondo and Takashi Oguchi ………………………………………………………056
Presentation of the PaleoGeo database with Google Earth
Stephen P. Obrochta, Yusuke Yokoyama and Hodaka Kawahata …………………………………………059
Regional last glacial climate synthesis for the Eastern Mediterranean
Ryouta O'ishi,Wing-Le Chan and Ayako Abe-Ouchi ………………………………………………………060
Vegetation change in Europe during the Late Pleistocene dispersal of Modern Humans
Yasuhisa Kondo ……………………………………………………………………………………………………061
Cost surface analysis based on ecological niche probability to estimate relative rapidity of the dispersals ofearly Modern Humans

Timing of the RNMH in Europe
Chair: Olaf Jöris

11:40-12:05

12:05-12:30
12:30-13:30
13:30-14:00
14:00-14:25

14:25-14:50

Katerina Douka and Tom Higham ……………………………………………………………………………064
Introduction to the “PalaeoChron” Project: Chronological mapping of the Middle and early Upper
Palaeolithic of Eurasia
Tom Higham ………………………………………………………………………………………………………065
Radiocarbon dating the final Mousterian of Eurasia using advanced chemistry methods
Lunch
Poster session
Takayuki Omori and Katsuhiro Sano …………………………………………………………………………066
Spatiotemporal pattern of the Middle to Upper Paleolithic transitional lithic industries in Europe: A new
approach for reliability assessment of radiocarbon data
Katsuhiro Sano and Takayuki Omori …………………………………………………………………………067
Replacement process of Neanderthals by Modern Humans in Europe

Global Processes of RNMH
Chair: Fereidoun Biglari

14:50-15:15

15:15-15:40

15:40-16:00
16:00-16:25

16:25-16:50

16:50-17:15
18:00

Seiji Kadowaki ……………………………………………………………………………………………………068
Diffusion or progressive integration? Re-considering the widespread distributions of early UP/LSA
bladelet technologies from west Asia to south Europe and from east Africa to south Asia
Paolo Biagi and Elisabetta Starnini ……………………………………………………………………………071
Neanderthals and Modern Humans in the Indus Valley (Sindh, Pakistan): The Middle and Late
Palaeolithic of a forgotten region of the Indian Subcontinent
Coffee break
Yoshihiro Nishiaki, Otabek Aripdjanov, Rustam Suleymanov, Hiroto Nakata, Saiji Arai,
Takehiro Miki, Takayuki Omori and Jannat Ismailova ………………………………………………………073
New insight into the end of the Middle Palaeolithic in Central Asia
Ekaterina A. Lipnina, German I. Medvedev, Fedora Khenzkhenova, Kunio Yoshida, Dai Kunikida,
Takao Sato, Kenji Suzuki and Hirofumi Kato …………………………………………………………………075
The exploitation and adaptation in the Asian Arctic and Paleolithic in the Baikal Siberia
Naruya Saitou, Timothy A. Jinam and Hideaki Kanzawa-Kiriyama ………………………………………077
Inferring possible contacts between Denisovan and some ancestors of modern East Eurasians
Free bus service to the Toyako Manseikaku Hotel Lakeside Terrace

DAY 4: December 3 (Wednesday), 2014
9:30-17:00

Registration

Excursion (optional)
10:00
Toyako Manseikaku Hotel Lakeside Terrace
10:10
Volcano Science Museum
12:00
Lunch at Shiraoi
13:00
Ainu Museum Poroto Kotan
17:00
Toyako Manseikaku Hotel Lakeside Terrace011

DAY 5: December 4 (Thursday), 2014
9:30-17:00

Registration

Learning Strategies and Cultural Evolution – Archaeological Approach
Chair: Nigel Goring-Morris

10:00-10:25
10:25-10:50
10:50-11:15

11:15-11:40
11:40-12:05
12:05-13:30
13:00-13:30

Olaf Jöris ……………………………………………………………………………………………………………080
Lives to learn? The biology and culture of Neanderthal and early Modern Human learning behaviours
Anna Belfer-Cohen and Erella Hovers …………………………………………………………………………081
TTT: Tradition, (archaeological) taxonomy and (cultural) transmission
Gilbert B. Tostevin ………………………………………………………………………………………………082
Cultural transmission in the Pleistocene: Studying Neanderthal-Modern Human contact in Central
Europe during OIS3
Foni Le Brun-Ricalens ……………………………………………………………………………………………084
Transculturation versus acculturation: A clarification
Jun Takakura ………………………………………………………………………………………………………085
Toward an understanding of learning strategy in the Upper Paleolithic: A case study in Hokkaido, Northern Japan
Lunch
Poster session

Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches
Chair: Hideaki Terashima

13:30-13:55

13:55-14:20
14:20-14:45

14:45-15:10

15:10-15:35
15:35-15:55

Bonnie Hewlett ……………………………………………………………………………………………………088
Innovation, processes of social learning and modes of cultural transmission amongst the Chabu
adolescent forager-farmers of Ethiopia
Barry S. Hewlett ……………………………………………………………………………………………………089
Cognition and cultural niche construction among Neanderthals and Modern Humans
Nobutaka Kamei …………………………………………………………………………………………………090
Theoretical and ethnographic studies on play and learning behaviors: The case of hunter-gatherer children in
the forest
Takeshi Furuichi, Crickette Sanz, Kathelijne Koops, Tetsuya Sakamaki, Heungjin Ryu,
Nahoko Tokuyama and David Morgan ………………………………………………………………………093
Why do wild bonobos not use tools like chimpanzees do? Examination of factors influencing the tool behaviors
Magnus Enquist, Stefano Ghirlanda and Johan Lind ………………………………………………………095
Some differences between humans and other animals
Coffee break

Learning Strategies and Cultural Evolution – Perspectives from Theoretical Modelling
Chair: Kenichi Aoki

15:55-16:20
16:20-16:45
16:45-17:10
17:10-17:35

012

18:00

Laurel Fogarty, Kenichi Aoki, Joe Yuichiro Wakano and Marcus Feldman ………………………………096
Population change and cultural variation
Shiro Horiuchi ……………………………………………………………………………………………………097
Why did the arts precede prehistoric settlement in Modern Humans?
Kenichi Aoki ………………………………………………………………………………………………………099
Population and culture: A theoretical basis for dramatic regime shifts?
Joe Yuichiro Wakano ………………………………………………………………………………………………102
Learning schedule and cultural evolution: COS and ESS
Free bus service to the Toyako Manseikaku Hotel Lakeside Terrace

DAY 6: December 5 (Thursday), 2014
9:30-17:00

Registration

Cognition and Learning
Chair: Hiroki C. Tanabe

10:00-10:25
10:25-10:50

10:50-11:15

Hideaki Terashima ………………………………………………………………………………………………106
The beginning of symbolic art and the learning cycle in nature
Keiichi Omura ……………………………………………………………………………………………………108
The ability of objectification as the basis for cultural learning and creativity: Teasing as the device fortraining children in Canadian Inuit society
Motoaki Sugiura, Sugiko Hanawa, Takayuki Nozawa, Rui Nouchi, Ryoichi Yokoyama, Yuka Kotozaki,
Tsuyoshi Araki and Ryuta Kawashima …………………………………………………………………………109
Neural correlates of coping strategy for boredom: Its relationship with creativity

Looking into the Neanderthal Brain
Chair: Naomichi Ogihara

11:15-11:40
11:40-12:05
12:05-13:10
13:10-13:35
13:35-14:00

14:00-14:25

14:25-14:50

14:50-15:10

Emiliano Bruner ……………………………………………………………………………………………………112
The evolution of the parietal areas in Neanderthals and Modern Humans: From bones to mind
Yasushi Kobayashi, Toshiyasu Matsui and Shigeru Kosuda …………………………………………………113
Estimating the extent of brain subdivisions using skulls
Lunch
Marcia S. Ponce de León and Christoph P. E. Zollikofer ……………………………………………………115
Inferring fossil brains from endocasts: New methods and results
Daisuke Kubo, Hiroki C. Tanabe, Osamu Kondo, Hideki Amano, Akira Yogi, Sadayuki Murayama,
Hajime Ishida and Naomichi Ogihara …………………………………………………………………………116
Estimating the cerebral and cerebellar volumes of Neanderthals and Middle andUpper Paleolithic Homo sapiens
Naomichi Ogihara, Hideki Amano, Yusuke Morita, Osamu Kondo, Daisuke Kubo, Hiromasa Suzuki,
Takashi Michikawa, Masaki Moriguchi, Hajime Ishida and Masato Nakatsukasa ………………………119
Endocasts of Neanderthals and early Modern Humans: Virtual reconstruction andgeometric morphometric analyses
Hiroki C. Tanabe, Takanori Kochiyama, Norihiro Sadato and Naomichi Ogihara ……………………121
Exploring the difference of brain anatomy and function between Neanderthals and Modern Humans:
Neuroanatomical and functional neuroimaging approach
Coffee break

Closing Remarks

Chair: Seiji Kadowaki

15:10-15:35
15:35-16:10
16:10-17:30
18:00

Yoshihiro Nishiaki …………………………………………………………………………………………………124
The RNMH project: A summary
Ofer Bar-Yosef ……………………………………………………………………………………………………125
Neanderthals and Modern Humans: The contributions of the RNMH project
Farewell party
Free bus service to the Toyako Manseikaku Hotel Lakeside Terrace

013

DAY 7: December 6 (Saturday), 2014
Post-conference Excursion (optional)
9:00
Toyako Manseikaku Hotel Lakeside Terrace
11:30
Lunch at Nibutani
12:30
Shigeru Kayano Memorial Museum, Saru Historical Museum and Ainu Cultural Museum
Lecture by Jun Takakura, Hokkaido University: Prehistory of the Hokkaido Island
16:00
New Chitose Airport
17:00
Sapporo JR Station

Poster Presentations (DAY 3 and DAY 5)
P1
P2
P3
P4
P5

P6
P7
P8
P9
P10

P11
P12

P13

014

Otabek Aripdjanov ………………………………………………………………………………………………128
Past and present research on the Middle and Upper Paleolithic of Uzbekistan
Yoshihiro Nishiaki, Yousef Kanjou and Takeru Akazawa ……………………………………………………131
The Neanderthal occupations at Dederiyeh Cave, Syria
Hiroto Nakata ……………………………………………………………………………………………………133
The emergence of the Upper Palaeolithic in the Japanese archipelago
Yasuo Higurashi …………………………………………………………………………………………………135
On the mechanics of the spear thrower
Eiko Yamagami ……………………………………………………………………………………………………137
Mediated action through art works among children: Similarities and differences between traditional andmodern society
Tadashi Koyama …………………………………………………………………………………………………139
Cognitive development of Baka Pygmy children growing up in modern hunter-gatherer society
Kaoru Imamura and Hiroyuki Akiyama ………………………………………………………………………142
Learning process on the skills of reading nature
Izumi Hagino and Taro Yamauchi ……………………………………………………………………………143
Fertility and population dynamics of Baka hunter-gatherers in southeast Cameroon
Ryosuke Kimura, Wataru Nakahashi and Kohei Tamura …………………………………………………145
The formation of cooperative society initiates the evolution of risk-taking innovative behaviors
Kyoko Yamaguchi, Toshiyuki Hayakawa, Chiaki Watanabe, Takehiro Sato, Akira Kawaguchi, ………148
Ken Yamamoto, Hajime Ishida and Ryosuke Kimura
Genome-wide association analysis of cephalic form in modern Japanese
Wataru Nakahashi ………………………………………………………………………………………………151
The effect of trauma on cultural transmission of Neanderthals
Yutaka Kobayashi, Seiji Kadowaki and Masaki Naganuma …………………………………………………154
A population-genetics based model for explaining apparent cultural continuity from the Middle toUpper Palaeolithic in Eurasia
Minoru Yoneda, Ayako Abe-Ouchi, Hodaga Kawahata, Yusuke Yokoyama and Takashi Oguchi ……156
Testing the hypothesis on cognitive evolution of Modern Humans’ learning ability: Achivements ofpast-climatic approaches

P14

P15

P16

P17
P18
P19
P20
P21
P22

Wing-Le Chan, Ayako Abe-Ouchi, Ryouta O’ishi and Kunio Takahashi ………………………………157
Stadial and interstadial climates in the Late Pleistocene, as simulated in experiments with the MIROC
climate modelFrancisco J. Jimenez-Espejo, Guadalupe Monge Gómez, Miguel Cortés Sanchez, Clive Finlayson,
Joaquin Rodriguez-Vidal and Antonio García-Alix …………………………………………………………159
Geochemistry of Neanderthal and Modern Human archeological sites: Evidences of paleoclimatic
variations and Homo s. activities
Takashi Michikawa, Masaki Moriguchi, Hiromasa Suzuki, Naomichi Ogihara, Osamu Kondo andYasushi Kobayashi …………………………………………………………………………………………………161
Endocast surface reconstruction from CT images
Osamu Kondo, Daisuke Kubo, Hiromasa Suzuki and Naomichi Ogihara ………………………………164
Reconstruction of Qafzeh 9 braincase and its endocast morphology
Masaki Moriguchi, Hiromasa Suzuki, Takashi Michikawa, Naomichi Ogihara and Osamu Kondo …166
Surface deformation of crania by optimizing rigidity
Naoki Miura, Yukako Sasaki, Kunihiro Hasegawa and Hiroki C. Tanabe ………………………………168
Neural underpinning of creative thinking: A meta-analysis for neuroimaging data
Yukinobu Hoshino, Keita Mitani, Naoki Miura, Hiroki C. Tanabe and Kenji Nagai …………………171
fMRI experiment about executive function by difference of learning methods: Using Tower of Hanoi
Kei Mizuno …………………………………………………………………………………………………………174
Brain structural and functional analyses for motivation to learn, fatigue and cognitive function
Kunihiro Hasegawa, Takanori Kochiyama, Hiroki C. Tanabe and Naomichi Ogihara …………………176
Reconstruction of the brain from skull fossil using computational anatomy

015

DAY 1: November 30, 2014

Opening Remarks
Keynote Lectures

Will we Modern Humans fade away or be replaced?

Takeru Akazawa

Research Institute, Kochi University of Technology, Japan. Email: akazawa0823@qd6.so-net.ne.jp

We humans, what are we? Various approaches to answering this question come to mind. One possibility is toinvestigate, through the numerous fossils of now-extinct hominids, how those hominids lived their lives. Thisshould provide some understanding of the path by which we have come to be who we are today. Among thosefossil hominids, there is much to be learned from the Neanderthals in particular, since they were the last hominid
population on Earth before Modern Humans took over.
Modern Humans appeared in Africa about 200,000 years ago. Later, as they spread across Eurasia, theyencountered indigenous Neanderthal populations. The two populations coexisted until 30,000 years ago, perhaps
even later, but the Neanderthals eventually went extinct. The question has long occupied us: what happened to the
Neanderthals? A number of current hypotheses propose possible mechanics of the replacement of Neanderthals
by Modern Humans, and there has been extensive debate as to whether or not the presence of the Modern
Humans accelerated the extinction of the Neanderthals.
Whenever human evolution reached such a fork, a new instance of humanity thrived while an old instancebecame extinct. In the last of these recurring succession dramas, the humans closest to us today, the modern Homosapiens, thrived while the earlier humans, archaic H. sapiens met their demise. All humans currently inhabiting
the earth are descendants of the anatomically modern H. sapiens who survived that epic transformation; now it isentirely possible that they are in midcourse towards the next succession event.
We can imagine the disappearance of today’s Modern Humans and their replacement by some new branch of
the human family, but no one knows what that new branch might be, nor can we concretely envision the natureof that new succession event. One long-term goal of humans, an abstract one, is to come to an understandingof the varied forms of existence we have experienced during our evolution. This quest may be of monumental
importance: essential clues to an understanding of our future may lie hidden in our past.

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Opening Remarks

From Neandertals to Modern Humans: A view from the
Italic Peninsula
Silvana Condemi

Anthropologie Bioculturelle, Droit, Ethique et Santé, Université d'Aix-Marseille/ CNRS/ EFS, Faculté de Médecine, France.
Email: silvana.condemi@univ-amu.fr

As in other regions of Europe, the presence of the Neandertal population on the Italian peninsula has been well
documented for a long time. The first evidence of Mousterian assemblages that, in Europe, were considered
to have been made solely by Neandertals, was identified at the end of the 19th century (in 1889) at the siteof Fate on the Italian riviera. The first Neandertal human fossil discovery occurred fortuitously in 1929, when
the nearly complete cranium of Saccopastore 1 was unearthed in the city of Rome. During the following
decade, this first discovery was augmented by that of two other major Italian fossils: Guattari 1, found in 1935
at the promotorium of Circeo (90 km south of Rome), and Saccopastore 2, unearthed in 1939 in the city ofRome. Although, once again, these two Neandertal crania were discovered by chance, and they lack of precise
stratigraphic context, they have contributed to our understanding of the Neandertals of southern Europe.
After a long interruption of research and exploration during the Second World War and the decade that
followed, excavations were begun throughout the central, northern and southern regions of the Italian peninsula
after the middle of the 1950s. At present, only a few excavations related to the middle and upper Pleistocene arestill in progress.
All research and study of the fossils and material uncovered by these excavations have permitted us to establish asimilarity in the history of the Italian Neandertals to that in other parts of Europe: after having lived and thrived
in Europe from MIS7 to MIS3, Neandertals then disappeared, leaving only Homo sapiens.
Although no verifiable Homo sapiens fossils from MIS3 have been found in Europe, their arrival is assumed dueto the presence of Aurignacian assemblages, the most ancient of which have been identified in eastern Europe.
From east to west, the dispersal of Homo sapiens resulted in the replacement of the local Mousterian Neandertal
population. In some cases, the Neandertal population was pushed toward marginal geographical areas, notably
toward the Iberian and Italian peninsulas.
During this process that occurred over a period of several thousand years, it was believed that the influence of
Homo sapiens in Europe led some Neandertal groups to modify their behavior and their way of life (d'Errico
et al. 1999) and, in particular, to change their Mousterian assemblages and produce what is often termed“transitional”assemblages (Szeletian, Chatelperronian, Uluzzian ... ). The assumption that Neandertals produced
all “transitional assemblages” was based on the well-preserved Neandertal remains at Saint Césaire that were
believed to have been associated with the Chatelperronian in southwestern France.
The hypothesis of a long period of Neandertal survival in marginal areas has found support in Spain due to thedates of Mousterian Neandertal sites (Finlayson et al. 2006) and, on the Italian peninsula, in view of the presenceof a “transitional”, Uluzzian assemblage limited to the central and southern parts of the peninsula. The presenceof Uluzzian culture (characterized by personal ornaments, bone tools and dyes) solely in the central and southern
regions of the Italian peninsula reinforced the impression that Neandertals who were pushed out found refuge inthis geographic area, where they continued to survive and began to make a “new” kind of assemblage influenced
by the technology of the incoming Homo sapiens. Indeed, according to this scenario, the Uluzzian assemblages

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Keynote Lecturesfound on the Italian and Balkan peninsulas were considered to be comparable to the Chatelperronian assemblages
found in France and in northern Spain. Since it was believed that the Chatelperronian was produced by late
Neandertals, it was assumed that Neandertals also made Uluzzian assemblages (Churchill et al. 2000).
In recent years this scenario has been completely overturned. First, the discovery of Uluzzian assemblages also
on the northern Italian peninsula, at the site of Fumane (Peresani 2008) shows that this kind of assemblage wasless confined to the south than had been supposed. Second, the new AMS dates of a Mousterian Neandertal site,
Riparo Mezzena near Fumane, indicate that Mousterian Neandertals may have survived longer than had beenthought (Longo et al. 2012).
Third and most important, however, is the re-analysis of the only human remains associated with Uluzzianarchaeological layers, two teeth from Grotta del Cavallo in the southern Italian peninsula found in 1964. In thisstudy (Benazzi et al. 2011), the teeth from Cavallo were compared by means of CT scans with those from a large
Modern Human and Neanderthal dental sample. The internal and external features of the dental crown, including
the thickness of the enamel and the general outline of the crown, were submitted to examination. The results
showed that the human teeth from Grotta del Cavallo were Modern Humans, not Neanderthals as had originallybeen thought. Furthermore, since the teeth were too small and precious to date directly, chronometric analysis
was carried out on marine shells found in the same archaeological levels as the teeth. This approach showed thedate of the Modern Human teeth to be between ~43,000-45,000 years ago (cal BP). This means that the humanremains are older than any other Modern Humans discovered in Europe and that these two teeth from Grottadel Cavallo therefore represent the oldest European Modern Human fossils currently known. The study of theteeth of Grotta del Cavallo confirms that the arrival of our species on the European continent, and the periodof their coexistence with Neanderthals, was several thousand years longer than was previously thought. Basedon this fossil evidence, this study also confirms that Modern Humans and not Neanderthals were the makers of
the Uluzzian culture and that the colonization of the European continent by Homo sapiens occurred in successive
waves.
Each of these three points is currently a topic of debate, which has been fueled by criticism regarding thechronostratigraphy of the Italian sites. The aim of our paper is to present the complex archeological data
originating from the Italian peninsula and to discuss problems and criticism which current studies of these sites
are likely to face.
References
Benazzi S et al (2011) Early dispersal of Modern Humans in Europe and implications for Neanderthal behavior. Nature 479:
525-528.
Churchill SE, Smith FH (2000) Makers of the Early Aurignacian of Europe. Yearbook of American Journal of Physical
Anthropology 43: 61-115.
d'Errico F et al (1998) Neanderthal acculturation in western Europe? Current Anthropology 39: 2–44.
Finlayson JC et al (2006) Late survival of Neanderthals at the southernmost extreme of Europe? Nature 443: 850-853.
Longo L et al (2012) Did Neandertals and anatomically Modern Humans coexist in northern Italy during the late Oxigen
Isotope Stage 3? Quaternary International 259: 102-112.
Peresani M (2008) A new cultural frontier for the last Neanderthals: The Uluzzian in northern Italy. Current Anthropology
49(4): 725-731.

020
Keynote Lectures

The expert performance model of Neandertal cognition

Thomas Wynn1* and Frederick L. Coolidge1

1. Center for Cognitive Archaeology, University of Colorado, USA. *Email: twynn@uccs.edu

With recent archaeological discoveries of Neandertal ornaments (Zilhao et al. 2010), the time-worn argumentthat Neandertals lacked symbolic culture is effectively defunct. Nevertheless, despite the many similarities
between the archaeological signatures of Neandertals and same-age Modern Humans, there are still apparent
differences. Some of these differences, such as Neandertals’ lower levels of innovation, suggest that there was amodest but significant cognitive difference between Neandertals and same-age Modern Humans (contra Villa
and Roebroeks 2014).
One way to understand this difference is to apply a formal model of cognition. For several decades cognitive
psychologists have studied a domain of thinking termed ‘expert performance’ which comes to the fore in welllearned, narrowly circumscribed activities such as chess and music. Several formal models have been proposed,
all of them focused on an expert’s ability to rapidly access large amounts of information held in long-term
memory. The specific model we have chosen to use is that of Anders Ericsson (K. Ericsson and Delaney 1999; K.
A. Ericsson and Kintsch 1995), largely because it incorporates working memory, a well-known cognitive ability
that the two of us have applied effectively to the study of recent cognitive evolution (Coolidge and Wynn 2005;
Wynn and Coolidge 2004). Ericsson’s model revolves around ‘retrieval structures,’ which are sets of cues held
in working memory that are linked to much larger encodings held in long-term memory (as when the retrieval
structure ‘King’s Indian Defense’ has cues linking to the locations of 32 chess pieces on the 64 square board forseveral successive moves). These retrieval structures are largely non-symbolic, and are learned by practice andrepetition, i.e., apprenticeship. Coincidentally, Keller and Keller (1996) have documented the kind of thinking
used by artisan blacksmiths, and their description is very similar to Ericsson’s account of expertise. When Keller
and Keller’s ethnographic account of technical cognition is combined with Ericsson’s psychological accountof expert cognition, the result is a powerful model of expert technical cognition that is directly applicable tothe archaeological record. Using this model we are able to account for many of the similarities and differences
presented by the archaeological signatures of Neandertals and same-age Modern Humans. In brief, Neandertals
appear to have relied very heavily on expert cognition as the basis for not just their technology, but also theirhunting and gathering.
This “Ericsson-Keller” model of technical cognition helps us understand the apparent difference in innovation.
Expert cognition powers flexible solutions to the large range of problems encountered in its narrow domain, but
it does not itself generate new solutions. If, as we suspect, Neandertals relied very heavily on expert cognition, we
would not expect to see much innovation, and we do not. Creativity relies on other cognitive systems, including
working memory capacity and, for technology, abstract concepts of folk physics. These are abilities that wereapparent for moderns, but not for Neandertals. Recent research on the role of the cerebellum in cognition
suggests that Neandertals’ comparatively smaller cerebellum might account for some of this lower level ofcreativity,
Finally, this Ericsson-Keller model is entirely consistent with the learning hypothesis as developed by Akazawa
and colleagues (Akazawa et al. eds. 2013, 2014). The kinds of learning required for expert performance are quite
different from the kinds of learning required for elaborate symbol-based, declarative information. Observation

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Keynote Lecturesand practice are the essential components, and language plays a much lesser role.
References
Akazawa T, Nishiaki Y, Aoki K (eds) (2013) Dynamics of learning in Neanderthals and Modern Humans vol. 1: Cultural
perspectives. Springer, New York.
Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) (2014) Dynamics of learning in Neanderthals and Modern Humans
vol. 2: Cognitive and physical perspectives. Springer, New York.
Coolidge FL, Wynn T (2005) Working memory, its executive functions, and the emergence of modern thinking. Cambridge
Archaeological Journal 15(1): 5-26.
Ericsson K, Delaney P (1999) Long-term working memory as an alternative to capacity models of working memory ineveryday skilled performance. In: Miyake A, Shah P (eds), Models of working memory: Mechanisms of active maintenanceand executive control. Cambridge University Press, Cambridge, pp. 257-297.
Ericsson KA, Kintsch W (1995) Long-term working memory. Psychological Review 102(2): 211-245.
Keller C, Keller J (1996) Cognition and tool use: The blacksmith at work. Cambridge University Press, Cambridge.
Villa P, Roebroeks W (2014) Neandertal demise: An archaeological analysis of the Modern Human superiority complex.
PLoS One. doi:DOI: 10.1371/journal.pone.0096424
Wynn T, Coolidge FL (2004) The expert Neandertal mind. Journal of Human Evolution 46: 467-487.
Zilhao J, Angelucci D, Badal-Garcia E, d'Errico F, Daniel F, Dayet L et al (2010) Symbolic use of marine shells and mineralpigments by Iberian Neandertals. Proceedings of the National Academy of Sciences 107(3): 1023-1028.

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Keynote Lectures

Neandertal: Abilities and realisations

Marcel Otte

Université de Liège, Belgium. Email: marcel.otte@ulg.ac.be

In ever aspects of behavior, Neandertal has proven his abilities. The Levallois is the most powerful everinvented in lithic techniques. The hunting strategy was highly elaborated. Burial pratices show their
metaphysic.
The only difference with Modern Man lies in their choices in the ways of life. These ways are in the context
of history, not anymore in the field of physical capacity. They choose the way of life that has been working
during 300,000 years.

023
Keynote Lectures

DAY 2: December 1, 2014

Neanderthals and Modern Humans: Behavioral Characterization
The First RNMH Processes in West Asia

The development of technology during the Earlier Stone Age:
Research at Wonderwerk Cave and the Kathu Complex, Northern
Cape Province, South Africa
Michael Chazan1*, Francesco Berna2, Steven Walker3, Ian Watts4,
Jayne Wilkins5 and Liora Kolska Horwitz6

1. University of Toronto, Canada; 2. Simon Fraser University, Canada; 3. University of Cape Town, South Africa; 4. Independent Scholar,
Greece; 5. Arizona State University, USA; 6. Hebrew University, Jerusalem, Israel. *Email: mchazan@chass.utoronto.ca

In the evolution of the genus Homo we cannot assume that technology is a passive expression of the cognitive
capacity, or invariable behavioural repertoire, of hominin populations (Chazan 2012). Members of genus
Homo display a capacity for technological innovation independent of speciation. Thus the relationship between
technological innovation and speciation is a significant issue in the study of the evolution of genus Homo. In
this paper I present an overview of recent research at Wonderwerk Cave and the sites of the Kathu Complex,
located in the Northern Cape Province, South Africa (Fig. 1). The results of this research point to a range oftechnological innovations in the time range between 1.8 mya and 0.5 mya. This is a period that can be correlatedwith H. erectus in a broad definition of this species that includes H. ergaster. The latter end of this period
corresponds with the initial appearance of the poorly defined group of hominins often described as archaic H.
sapiens. The results emerging from research at Wonderwerk Cave and the sites of the Kathu Complex suggestthat technological innovation played an active role in the dynamics of the evolution during this time period.
Wonderwerk Cave penetrates 140 meters into the dolomitic rocks on the eastern flank of the Kuruman Hills.
Excavations by Peter Beaumont (Beaumont and Vogel 2006) exposed a long sequence of Earlier Stone Age (ESA)
occupation in the front of the cave (Excavation 1) and an occupation attributed to the transitional (Earlier Stone
Age/Middle Stone Age) Fauresmith industry at the very back of the cave (Excavation 6) (Fig. 2). Our researchteam has undertaken a program of study to develop an absolute chronology for Excavation 1 along with acharacterization of lithic and faunal material recovered by Beaumont. In December 2013 we initiated a programof excavation at the site which will expand the excavated area and allow for the recovery of high precision spatial
data. A chronology for the lower levels of Excavation 1 has been developed on the basis of paleomagnetic
and cosmogenic burial age dating (Chazan et al. 2008, 2012; Matmon et al. 2012; Ruther et al. 2009) (Fig 2).
The earliest level (St. 12), dating to the Olduvai event (1.96-1.78 mya) contains a small stone tool assemblage
attributed to the Oldowan (Chazan et al. 2012). The first bifaces appear in St. 11 dated to 1.78-1.07 mya. In St.
10, dated to the Jarmaillo (1.07- 0.99 mya), handaxes are the dominant component of the assemblage. Handaxes
in St. 10 and in overlying St. 9 do not show shaping through invasive flake removals. Highly symmetricalhandaxes shaped by invasive removals first appear in St. 8 and continue to dominate the assemblage through theend of the ESA sequence in St. 5. Unfortunately, St. 9-5 have not yet been dated although this is a major goalof the new program of excavation. Cleavers are found beginning is St. 10 and it is particularly interesting that acleaver made with the Victoria West method is found in St. 9.
In addition to the evidence for developments in lithic technology, Excavation 1 has yielded evidence for theearliest known presence of fire in an in situ archaeological context. Based on a combination of micromorphology,
FTIR, and macroscopic evidence on lithic and faunal remains there is clear evidence for burning in St. 10
dated to the Jaramillo (Berna et al. 2012). There is limited evidence for burning in St. 12 based on analysis of
microfauna (Chazan et al. 2012).

026

The back of Wonderwerk Cave, Excavation 6, is poorly constrained chronologically but can be securely attributed
to the Fauresmith transitional industry. The lithic component includes bifaces, blade production, and prepared

Neanderthals and Modern Humans: Behavioral Characterizationcore technology. The position of Excavation 6, 140 meters from the front of the cave, and the character of thearchaeological assemblage which includes in addition to stone tools non-utilitarian objects such as quartz crystals
and specularite have been interpreted by Chazan and Horwitz (2010) as evidence of hominin sensitivity to thesensory properties of places and object that is the precursor to symbolic behaviour.
The sites of the Kathu Complex, located on the western flank of the Kuruman Hills, provide an opportunity to
explore the Acheulean and Fauresmith in the context of high density open air sites. At the Kathu Townlandssite there is a remarkable density of artifacts attributed to the Acheulean that raises questions about the natureof hominin mobility and group size (Walker et al. 2014). At the Bestwood 1 site a horizon of Fauresmith
occupation is found buried below sands covering hectares providing the potential to explore hominin organizationof activities across the landscape (Chazan and Wilkins et al. 2012). At Kathu Pan 1, Acheulean and Fauresmithmaterial is found in a complex stratigraphic context effecting by the action of spring vents. The Fauresmith
Stratum 4a has been dated by a combination of OSL and ESR/U-series to ca. 500kyr (Porat et al. 2009). The
lithic assemblage from Stratum 4a provides early evidence of blade production and the use of stone tipped spears(Wilkins and Chazan 2012; Wilkins et al. 2012).
References
Beaumont PB, Vogel JC (2006) On a timescale for the past million years of human history in central South Africa. South African
Journal of Science 102 (5-6): 217.
Berna F, Goldberg P, Horwitz LK, Brink J, Holt S, Bamford M, Chazan M (2012) Microstratigraphic evidence of in situ fire in the
Acheulean strata of Wonderwerk Cave, Northern Cape Province, South Africa. Proceedings of the National Academy of Sciences
109: E1215-E1220.
Chazan M (2012) Handaxes, concepts, and teaching. Mind, Brain, and Education 6(4): 197-203.
Chazan M, Avery MD, Bamford MK, Berna F, Brink J, Fernandez-Jalvo Y, Goldberg P, Matmon A, Porat N, Ron H, Rossouw
L, Scott L, Horwitz LK (2012) Oldowan occupation at Wonderwerk Cave, South Africa: Archaeology, geology, paleontology, andpaleoclimate. Journal of Human Evolution 63: 859-866.
Chazan M, Horwitz LK (2010) Milestones in the development of symbolic behaviour: A case study from Wonderwerk Cave, South
Africa. World Archaeology 41(4): 521-539.
Chazan M, Ron H, Matmon A, Porat N, Goldberg P, Yates R, Avery M, Sumner A, Horwitz LK (2008) First radiometric dates forthe earlier stone age sequence in Wonderwerk Cave, South Africa. Journal of Human Evolution 55: 1-11.
Chazan M, Wilkins J, Morris D, Berna F (2012) Bestwood 1: A newly discovered Earlier Stone Age living surface near Kathu,
Northern Cape Province, South Africa. Antiquity 86(331): Antiquity Gallery.
Jacobson L, de Beer FC, Nshimirimana R, Horwitz LK, Chazan M (2012) Neutron tomographic assessment of incisions on
prehistoric stone slabs: A case study from Wonderwerk Cave, South Africa. Archaeometry 55(1): 1-13.
Matmon A, Ron H, Chazan M, Porat N, Horwitz LK (2012) Reconstruction the history of sediment deposition in caves: A casestudy from Wonderwerk Cave, South Africa. Geological Society of America Bulletin 124 (3-4): 611-625.
Porat N, Chazan M, Grun R, Aubert M, Eisenmann V, Kolska-Horwitz L (2009) New radiometric ages for the Fauresmith industry
from Kathu Pan, Southern Africa: Implications for the Earlier to Middle Stone Age Transition. Journal of Archaeological Science37: 269-283.
Riddle ATR, Chazan M (2014) Stone tools from the inside out: Radial point distribution. World Archaeology ahead-of-print: 1-14.
Ruther H, Chazan M, Schroeder R, Neeser R, Held C, Walker SJ, Matmon A, Horwitz LK (2009) Laser scanning for conservation
and research of African cultural heritage sites: The case study of Wonderwerk Cave, South Africa. Journal of Archeological Science
38: 1847-1856.
Walker, SJH, Lukich V, Chazan M (2014) Kathu Townlands: A high density Earlier Stone Age locality in the interior of South
Africa. PloS one 9.7: e103436.
Wilkins J, Schoville BJ, Brown KS, Chazan M (2012) Evidence for early hafted hunting technology. Science 338(6109): 942-946.
Wilkins J, Chazan M (2012) Blade production ~500 thousand years ago at Kathu Pan 1, South Africa: Support for a multiple origins
hypothesis for early Middle Pleistocene blade technology. Journal of Archaeological Science 39(6): 1883-1900.

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Neanderthals and Modern Humans: Behavioral Characterization

Fig. 1 Map showing location of Wonderwerk and the Kathu Complex. WW: Wonderwerk; BW:
Bestwood; KT: Kathu Townlands; KP: Kathu Pan.

Fig. 3 Stratigraphic profile of Excavation 1 Wonderwerk Caveshowing dating samples and location of micromorphologicalevidence of wood ash.

Fig. 2 Plan of Wonderwerk Cave (Curtesy of H. Ruther) showing
excavation areas.

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Fig. 4 Artifacts from the Bestwood 1 site.

Neanderthals and Modern Humans: Behavioral Characterization

A tale of three caves: The role of mammoths in Neanderthal
adaptation and lifeways
Ran Barkai

Department of Archaeology and Near Eastern Cultures, Tel-Aviv University, Israel. Email: barkaran@post.tau.ac.il“The elephant is the largest of them all, and in intelligence approaches the nearest to man” (Bostock, J., & Riley, H. T.,
1855. The Natural History. Pliny the Elder. Taylor and Francis, London)
The Paleolithic archeological record depicts complex relationship between humans and elephants, as elephants
were exploited for their meat, fat and bone over hundreds of thousands of years across the old world.
The role of protein in human diet and subsistence in Paleolithic sites has been demonstrated time and again.
Animal meat and fat are an excellent source for calories, and contain essential amino acids, minerals, vitamins andfatty acids.
As early as the late Pliocene we are familiar with archaeological sites containing stone tools associated with animals' bones, reflecting human activities related to butchery of animals and marrow extraction. Hence, it is commonly accepted that Acheulian, and even pre-Acheulian hominins, extracted a significant portion of the calories
they consumed from animal meat and fat, and thus were actually dependent on animals for their successful survival. Hence, carnivory is a remarkable human trait accompanying the human race from its earliest stage to thisvery day.
Palaeolithic nutrition was based on animal meat and fat in addition to plant-based foods. Many Paleolithic sites
have extensive evidence for large mammal consumption and it has been argued repeatedly that big game hunting
was a principal procurement strategy, most probably accompanied by the occasional use of carcasses resulted fromnatural death or leftovers by other high carnivores.
The significant role of Proboscideans in Paleolithic faunal assemblages is well demonstrated in many sites in Europe, Africa and Asia. However, the dietary significance of elephants has not been thoroughly explored. We work
under the assumption that during Paleolithic times Proboscideans, when available, were a constant and significant
source of calories for early hominins. Notwithstanding its significance, this was only one source of calories among
many other (plant based and animal based) food resources used by Paleolithic hominins. However, in our opinion,
it was a primary one when Proboscideans were available.
The use of elephant bones in the Acheulian for the production of tools, particularly bifaces that resemble thecharacteristic stone handaxes, is clearly evident. Several Lower Paleolithic Acheulian sites, however, clearly
demonstrate butchery and defleshing of elephants. Post-Acheulian as well as Mousterian sites provide further
evidence for the use of Proboscideans both for dietary purposes, such as meat and marrow consumption, and forother tasks, such as bone tool production.
Proboscideans were by far the largest terrestrial animal available for Lower Paleolithic hominins. They represent
an ideal food package with a perfect combination of meat and fat, as half of the potential calories are in the fat.
While we have no exact data as to the extent to which elephant carcasses were utilized, the abundance of evidence for elephant utilization in Paleolithic sites is a clear indication that at least part of the elephant’s potential

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Neanderthals and Modern Humans: Behavioral Characterization

energetic value was extracted by early humans. This view is further supported not only by the fact that some elephant bones found in Paleolithic sites bear cut marks, but also by the fact that many bones were further fractured
in order to reach the marrow, indicating further use of the bones for additional fat.
In this presentation I will present several case studies for the use of Proboscideans in Lower and Middle Paleolithic sites from the Old World, discuss the role of Elephants and Mammoths in early human adaptation, dietand culture and explore the significance of Mammoths in Neanderthal survival and life-ways. An attempt will bemade to correlate between the systematic use of Mammoths by European Neandethals and knowledge transmission mechanism that has to do with the acquisition and exploitation of such gigantic mammals.

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Neanderthals and Modern Humans: Behavioral Characterization

Middle Palaeolithic flint mines in Mount Carmel: An
alternative interpretation
Avraham Ronen

Zinman Institute of Archaeology, University of Haifa, Israel. Email: aronen@research.haifa.ac.il

The situation
Numerous heaps of limestone fragments around Nahal (river) Galim in north-western Mount Carmel wereinterpreted as residues of quarries. The benefit of the operation would be to extract flint nodules lying beneathlimestone beds and the lithic production areas. Freshly extracted flint nodules are indeed better knapped than
nodules collected on the surface, battered by the elements and desiccated under intense sunshine. Knapped flint
nodules were taken to indicate workshops of lithic production. A few Middle Palaeolithic flint artifacts mingled
in the limestone heaps suggested Middle Palaeolithic date for the quarries and the hypothesized knapping
workshops (Nadel et al. 2011).
Significance
From a technological perspective, experts of the Levallois method had no doubt the ability to break a limestone
bed. However, this hypothesis holds far reaching consequences on the social organization of Middle Palaeolithic
hominins in the Levant. The society would be able to assemble individuals for the tedious work of quarrying(perhaps even those not in immediate need of flint). A mechanism to mobilize individuals for a communal effort
reflects a social organization more advanced than generally assigned to Middle Palaeolithic hominins.
Difficulties
Two observations render, however, the quarry/workshop hypothesis problematic: first, the low density of lithic
products among the limestone heaps (Nadel et al. 2011), and second, the virtual absence of clearly delineatedconcentrations of flint knapping residues - nodules, cores, flakes and debris. Unquestioned knapping workshops(Neolithic, for example) consist of well defined concentrations of knapping residues (Ronen and Davies 1970;
Taute 1994).
Another interpretation
An alternative interpretation of the Mount Carmel limestone heaps is offered here. While admitting theanthropogenic origin of the heaps, this interpretation questions their function and the Middle Palaeolithic age
assigned to them. Accordingly, rather than Middle Palaeolithic quarrying debris, the heaps would constitute raw
material for the manufacture of lime during the last three millennia or so (Sasson 2002).
Lime was an important element in building the traditional Levantine house (Peled 2010). Building a housestarted by building a lime kiln at the site (Canaan 1933). Large quantities of lime were used: mixed with sand, thelime served as mortar to bind the building stones while mixed with clayey earth, it served as plaster to be applied
on walls and roof to repel atmospheric humidity (Canaan 1933). In the immediate area of the heaps discussed
here around Nahal Galim, remains of some 20 lime kilns were revealed by the Archaeological Survey of Mount
Carmel (Ronen and Olami 1978; Olami et al. 2003). Each kiln was fed ca. 4-5 tons of limestone fragments. The
stones were burnt during 4-5 days until they became lime. Then the kiln was left to cool for 3-5 days before thelime could be removed and used. Hence every 10 days on the average, some 100 tons of limestone fragments wererequired, i.e. approximately 10 tons per day, to feed the kilns in the research area.

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Neanderthals and Modern Humans: Behavioral Characterization

Conclusions
Middle Palaeolithic flint artifacts are abundantly scattered on the surface of Mount Carmel (Olami 1984). Someof these artifacts became naturally mingled in the limestone heaps. These intrusive finds do not denote the age ofthe heaps. The heaps are not of Middle Palaeolithic age, nor are they quarry debris. They were, in all likelihood,
raw material for lime production during historical times.
References
Canaan T (1933) The Palestinian Arab house: Its architecture and folklore. Journal of the Palestine Oriental Society 13: 1-83.
Nadel D, Rosenberg D, Shtober Zisu N, Filin S (2011) The Nahal Galim/Nahal Ornit prehistoric flint quarries in Mt. Carmel,
Israel. Eurasian Prehistory 8 (1–2): 51–66.
Olami Y (1984) Prehistoric Carmel. Israel Exploration Society, Jerusalem.
Olami Y, Ronen A, Romano A (2003) Map of Haifa-West. Israel Antiquities Authority, Jerusalem.
Peled K (2010) Architecture, the Arab house as a social text. Resling, Tel-Aviv.
Ronen A, Davies M (1970) Un atelier de taille néolithique au Mt. Carmel: Le Point z 355. L'anthropologie 74: 161-194.
Ronen A, Olami Y (1978) Atlit map. Archaeological Survey of Israel, Jerusalem.
Sasson A (2002) Lime kilns on the Carmel. Michmanim 16: 41*-42*.
Taute W (1994) Pre-Pottery Neolithic flint mining and flint workshop activities southwest of the Dead Sea, Israel (Ramat Tamarand Mesad Msazzal). In: Gebel HGK, Kozlowski SK (eds) Neolithic chipped lithic industries of the Fertile Crescent. Ex-oriente,
Berlin, pp. 495-510.

Fig. 2 A heap of limestone fragments in Mount Carmel (photo, A. Ronen).

Fig. 1 Location map of the study area (redsquare) among MP burial sites.

032
Neanderthals and Modern Humans: Behavioral Characterization

Fig. 3 Flint nodules knapped.

Fig. 4 Cross section of a lime kiln (from Sasson 2002).

Fig. 5 Remains of a lime kiln (from Olami et al. 2003).

Fig. 6 Area of the Nahal Galim stone heaps (circle) and location of lime kilns (squares).

033
Neanderthals and Modern Humans: Behavioral Characterization

A week (?) in the life of the Mousterian hunter

Gonen Sharon

Tel Hai College, Israel. Email: gonen@telhai.ac.il

Eight excavation seasons at the Mousterian site of Nahal Mahanyeem Outlet (NMO) on the banks of
the Upper Jordan River offer a glimpse into the life ways of MP people during a hunting expedition in
the Northern Dead Sea Rift. This open-air site, OSL dated to ca. 60ky BP, is interpreted as recording aseries of short-term hunting events. The NMO horizons, with their small number of lithic artifacts, uniquetypological composition and evidence for task specific hunting and butchering activity fit within Binford's(1980) definition of a “task location”. Many of the models suggested to describe site pattern and mobility
activity, such as the foraging and logistical models, are based primarily upon theoretical consideration andethnographic evidence. NMO gives us the opportunity to test such models based on archaeological evidence.
While still in dispute to some extent (the way archaeological questions always will be) the “large scale”
questions of the Middle Palaeolithic (MP) in the Levant are now defined and data is available upon which
to base answers. The chronological framework for the Levantine MP is established, the human biologicaland anatomical diversity is documented and we have a chronological sequence for the lithic phases of theLevantine MP. We can now turn to the higher resolution questions that NMO is uniquely suited to explore.
These crucial questions concern the life-ways of the people, their subsistence strategies and diet, functionalityof their tools, their specific tool kits and their hunting practices. An additional aspect of archaeologicalresearch that is in need of new data and study is that of the environmental background for human presence
during the Middle and Late Pleistocene, a period for which the actual Levantine climatic conditions are verymuch in debate.
NMO’s findings include a unique, small lithic assemblage comprising the highest percentage of tools to waste
recorded in the Levant. The assemblage is characterized by many pointed elements and cutting elements.
Importantly, the assemblage contains a very low percentage of other tool types such as scrapers. Refittingof several reduction sequences from the assemblage has enabled us to explore questions of lithic technology
and raw material economy. The NMO findings also include excellently preserved organic material, including
animal bones, both micro and macro-fauna and botanic remains (wood, seeds fruit and pollen).

034

NMO is dated to the late stage of the Levantine MP, contemporary to the presence of Neanderthal
populations in the region at sites such as the Amud Cave and Kebara Cave. While the probability of
exposing human remains at an open-air site like NMO is low, the site has great potential for exploring
crucial questions regarding behavior of MP hominins at the brink of the appearance of anatomically Modern
Humans. The small size of the assemblage, the unique typology of the artifacts and the extraordinarypreservation of the bones, wood, fruits and seeds at NMO are but one aspect of the site's potential. We
have now acquired information and data that enable us to suggest models that reconstruct the nature of
the site and human behavior reflected in its layers and finds. Issues of cognitive abilities, hunting praxis andsubsistence strategies, site function and group mobility all are in need of new, high resolution data that NMO
can provide. Moreover, the high resolution stratigraphy and, above all, the excellent preservation of botanicand pollen remains can achieve a comprehensive picture of the Late Pleistocene environment and its impact
on human diet, subsistence and survival.

Neanderthals and Modern Humans: Behavioral Characterization

Fig. 1 Location map of the NMO site.

Fig. 2 Refitted reduction sequence from Area D at NMO.

Fig. 3 Fauna of NMO: 1. Bovid skull compared to modern day
skull; 2. Skull and femur of lion (panther leo); 3. Wild boar
mandible; 4. Bos femur.

035

Neanderthals and Modern Humans: Behavioral Characterization

Evolution of learning capacity and acquired behaviors ofModern Humans from the perspective of life history andbrain development
Taro Yamauchi

Faculty of Health Sciences, Hokkaido University, Japan. Email: taroy@med.hokudai.ac.jp

Among primates, Modern Humans (Homo sapiens) have a unique growth pattern: a prolonged period ofchildhood and a rapid growth spurt in adolescence (Fig. 1). It is considered that the long childhood period
evolved in order to promote brain growth and development by depressing body growth rate. In reality, brain
growth is much faster than body growth and it completes at the ages of 7–10 (in weight). Once brain growthis complete, a rapid body growth (the so-called adolescent growth spurt) occurs to bring the body to adult
size (Fig. 1).
The stages of human life history after birth are as follows: infancy, childhood, juvenile, adolescence, and adulthood(Table 1). When we consider the growth rate (e.g., height [cm/year]) that takes place during each of these stages,changes in growth rate are clearly associated with each stage of development (Fig. 1). Each stage also may be
defined by characteristics of dentition, changes related to methods of feeding, physical and mental competencies,
and maturation of the reproductive system and sexual behavior.
The unique growth pattern of humans coincides with the development of cognitive ability, behavior, andmind. Moreover, we consider not only body growth but also the brain development to be the foundations oflearning capacity and acquired behaviors. Adolescence is a unique period for the body to grow, but also for brain
development. As previously mentioned, the brain reaches approximately 90% of its adult size by the age of six,
but the brain continues to undergo dynamic changes throughout adolescence.
The brain’s axons become gradually more insulated with myelin, eventually boosting the axons’ transmission speed
up to a hundred times. While myelin sheathing greatly accelerates an axon’s bandwidth, it also inhibits further
axonal branching. The process of maturation continues throughout adolescence and is completed by mid-20s. Thisdelayed completion heightens the flexibility of the brain and appears to be a unique human adaptation.
Adolescents lack consideration and engage in high-risk behaviors. Conventional brain studies have attributed thisphenomenon to immaturity of the brain. Recent research, however, considers adolescent behavior to be positivefrom an evolutionary perspective. The differences in learning capacity and acquired behavior as well as creativity
may be attributed to differences in brain development during the stages of adolescence between the Neanderthals
and Modern Humans.
In this presentation, I will propose some possible scenarios about the life history and the brain developmentof the Neanderthals to find a way to verify the “learning hypothesis” that the Neanderthal-Homo sapiens
replacement was the result of the difference in learning capacity between archaic and Modern humans.

036

References
Bogin B (1999) Pattern of human growth, 2nd edn. Cambridge University Press, Cambridge.
Leonard WR, Snodgrass JJ, Robertson ML (2012) Comparative and evolutionary perspectives on human brain growth. In: Cameron
N, Bogin B (eds) Human growth and development, 2nd edn. Academic Press, London.
Casey BJ, Jones RM, Hare TA (2008) The adolescent brain. Annals of the New York Academy of Sciences 1124: 111-126.
Baird AA, Fugelsang JA (2004) The emergence of consequential thought: Evidence from neuroscience. Philosophical Transactions of

Neanderthals and Modern Humans: Behavioral Characterization

the Royal Society of London B 359: 1797-1804.

Fig. 1 Human growth pattern (height) and life stages.
I: infancy; C: childhood; J: juvenile; A: adolescence; M:mature adult.

Table 1 Stages in the human life cycle.

Life stage
Birth
InfancyChildhood
Juvenile
Adolescence
Mature adult

Approximate ages (years old)
0
0-3
3-7
7-10 (boys), 7-12 (girls)
10-18 (boys), 12-18 (girls)
18+

037
Neanderthals and Modern Humans: Behavioral Characterization

The Early Middle Paleolithic of Misliya Cave, Mount Carmel,
IsraelMina Weinstein-Evron1*, Yossi Zaidner1, Alexander Tsatskin1,
Reuven Yeshurun1 and Israel Hershkovitz2

1. Zinman Institute of Archaeology, University of Haifa, Israel; 2. Department of Anatomy and Anthropology, Sackler Faculty of Medicine,
Tel-Aviv University, Israel. *Email: evron@research.haifa.ac.il

The long-collapsed Misliya Cave, Mount Carmel, Israel, located some 7 km north of Tabun Cave, contains
Lower Paleolithic (Acheulo-Yabrudian) and Early Middle Paleolithic (EMP, Mousterian) layers. It is one of
the rare Levantine sites in which both the Acheulo-Yabrudian and Early Mousterian are present, allowing
direct comparison between the two industries. The Acheulo-Yabrudian, TL dated to ~250 ka ago, was foundmainly on the lower terrace of the site. It is characterized by several technological systems including bifacial
shaping; production of thin flakes from unidirectional cores and cores with one preferential flaking surface
and prepared striking platform; production of large and thick, often cortical, flakes from unprepared cores.
The thick flakes were used for manufacturing of handaxes and of large scrapers by Quina or semi-Quina
retouch.
The EMP layers, spread on the upper terrace of the site, are between 1-3.5 m in depth and contain a series ofwell-defined hearths and an extremely rich lithic industry (Tabun D-type). TL dates on burnt flints suggest
dates of ~250-160 ka to EMP layers. The emergence of the EMP is manifested by a technological break,marked by the disappearance of bifaces and thick-flake production technology together with the introductionof blade manufacture using laminar and Levallois production methods, and the Levallois method for point
and triangular flake production. The tool-kit is dominated by various types of points and retouched blades.
No significant changes in material culture were observed along the EMP sequence. The repeated constructionof hearths, the abundant burnt flints and bones, and the high density of finds (ca. 3,000 lithic artifacts larger
than 2.5 cm per cubic meter) suggest intensive occupation at the site and attest to the habitual use of fire.
The rich faunal assemblage is overwhelmingly dominated by ungulate taxa with Mesopotamian fallow deer
(Dama mesopotamica) and mountain gazelle (Gazella gazella) being the most common prey species. Bodypart analysis, age profiles and bone-surface modifications observed at this almost exclusively anthropogenic
assemblage all attest to systematic ungulate hunting and ‘modern’ carcass transport and butchery patterns.
Hunting was performed using sophisticated gear, employing a large variety of points of different forms andsizes. Phytoliths and charred vegetal remains indicate extensive use of plants and exhibit the earliest use of
bedding to date. The spatial arrangement of EMP layers indicates that the site was used largely in the same
way throughout the occupation. Recurrence of the same spatial pattern throughout the EMP sequence,
together with high density of artifacts and bones and repeated use of fire suggest repeated visits and low
residential mobility, much like in the late Mousterian sites in the region.
The sharp differences between Acheulo-Yabrudian and EMP lithic technologies indicate that the transition
from the Lower to Middle Paleolithic in the Levant was rapid and may imply the arrival of a new population
around 250 ka. Human remains are scarce. Their study will shed important light on the emergence ofModern Humans.

038
The First RNMH Processes in West Asia

Preliminary study of the lithic assemblages from Nesher Ramla:
A case of continuity and change in the Levantine Middle
Paleolithic
Yossi Zaidner1,2*, Laura Centi2, Marion Prevost2, Maayan Shemer1,3 andOz Varoner4

1. Zinman Institute of Archaeology, University of Haifa, Israel; 2. Institute of Archaeology, the Hebrew University of Jerusalem, Israel; 3.
Israel Antiquities Authority, Israel; 4. Department of Bible, Archaeology and the Ancient Near East, Ben-Gurion University of the Negev,
Israel. *Email: yzaidner@research.haifa.ac.il

A recently discovered eight-meter-thick open-air sequence at Nesher Ramla (Israel) provides new evidence
regarding Levantine Middle Paleolithic adaptations during most of MIS 6 and 5 (OSL dates: 167±11 ‒
78±6 ka). The site is located in a karst depression formed by gravitational deformation and sagging into
underground voids. During the deposition of the archaeological sequence the formation of the site wasprimarily shaped by a cyclic mechanism of deposition of eroded soils, water-logging and pedogenesis.
Excavations at the site yielded rich lithic and faunal assemblages, combustion features, hundreds ofmanuports, hammerstones and anvils, and ochre. The eight-meter-thick archaeological sequence was divided
into six stratigraphic units, in which several horizons and vertically distinct concentrations of lithics andbones were identified.
The long archaeological sequence at Nesher Ramla allows testing some of the models on the Levantine
Middle Paleolithic technological variability, continuity and change. The 200,000 years long Levantine
Mousterian sequence show a little technological diversity and is dominated by Levallois reduction strategy.
During the early part of the Levantine Mousterian (250-160 ka), Levallois reduction strategy coexisted withsystematic use of laminar technology, but it became the dominated formal reduction strategy during the laterpart of the Mousterian (160-45 ka).
Variability within the Levallois reduction strategy has long been a subject for heated debate. Some scholars
see a shift from recurrent centripetal to recurrent convergent unipolar Levallois methods around 70 ka andsuggest that it was associated with the arrival of the Neanderthals. Others suggest that assemblages prevailed
by either of two methods are not arranged in the chronological order and rather represent group preferences
within a general population and cultural continuity in the Levant.
We will present a technological analysis of the lithic assemblages from main archaeological units of Nesher
Ramla and discuss their implication for the understanding of the technological continuity and change
between 170-80 ka, and especially for the shift from centripetal to convergent modes of Levallois flaking.

039
The First RNMH Processes in West Asia

Technological trends of the Middle to Upper Paleolithic
transition at Sefunim Cave, Israel
Andrew W. Kandel1* and Ron Shimelmitz2

1. The Role of Culture in Early Expansions of Humans (ROCEEH), Heidelberg Academy of Sciences and Humanities, University of
Tübingen, Germany; 2. Zinman Institute of Archaeology, University of Haifa, Israel. *Email: a.kandel@uni-tuebingen.deIn his monograph about the archaeology of Sefunim Cave in Israel, Ronen (1984) reported the sequence tobe transitional between the Middle and Upper Paleolithic. However, detailed technological studies of thelithic assemblages were lacking. Furthermore, studies about the large mammalian fauna, micromammals,
paleoecology and geoarchaeology were also needed to place technological developments at the site into an
ecological framework. Therefore, we established a joint mission between the Universities of Tübingen and
Haifa to re-examine this important Levantine sequence. The aims of this project are to collect new data
about unresolved issues by: 1) investigating behavioral patterns and the environment; 2) correlating changes
in behavioral patterns with fluctuations in the environment; 3) analyzing mobility patterns throughout
the sequence; 4) examining the possibility of population movements during the Upper Paleolithic and 5)
assessing whether evidence for acculturation exists.
Sefunim Cave (also known as Iraq el-Barud) is located 2.7 km east of the Mediterranean coastline in the
Carmel Mountain National Park (Fig. 1). The cave is situated at an altitude of 125 m above sea level in asteeply incised valley known as Nahal (Wadi) Sefunim. Located near the base of a massive limestone outcrop
and opening to the north, this cave complex consists of three adjoining chambers and additional niches alongits 60 m length and 20 m width. The front chamber contains some of the largest dripstone features known inMount Carmel.
Stekelis (1961) dug a test excavation in 1941 and uncovered a 1.2 m sequence with an Aurignacian horizon
at its base and overlying Kebaran, Natufian and Pre-Pottery Neolithic (PPN) layers. Ronen (1984) conductedspatially extensive excavations from 1965-1969 and succeeded in documenting a 2.0 m sequence startingin the Mousterian and continuing through Initial Upper Paleolithic, Levantine Aurignacian, Natufian and
PPN occupations (Table 1). He also discovered prehistoric occupation on the terrace outside of the cavewith Mousterian deposits overlain by PPN. Ronen pinpointed the location of the Mousterian layers to
the entrance and terrace of the cave and recognized that the Upper Paleolithic layers were more extensive
than the Middle Paleolithic horizons. He attributed large rockfalls at the entrance of the cave to Holocene
collapse, and these likely helped preserve the archaeological deposits.
In May 2013 the authors renewed field activities at Sefunim. The first season included two weeks offieldwork and confirmed the presence of intact stratified sediments. The excavation at the drip line covered
3 m2 and extended to a depth of 45 cm, while a single square meter dug on the terrace reached a depthof 55 cm. Excavation at the drip line encountered the top of Ronen’s Levantine Aurignacian layer 8 (Fig.
2). The new excavation unearthed sediments rich in finds, including numerous lithic artifacts and faunal
remains, as well as shell beads, bone tools and ocher. In contrast to the excavation at the drip line, the terrace
excavation yielded few finds, but the presence of Levallois technology allowed their attribution to the Middle
Paleolithic.

040

Preliminary studies of the lithic materials show that our layers II-IV are characterized primarily by blade
and bladelet production. Waste includes cores, core trimming elements, primary element flakes and regular

The First RNMH Processes in West Asia

flakes. Blank production was aimed at the manufacture of narrow and elongated bladelets characteristic of
the early Epi-Paleolithic. Among the microlithic tools, backed and obliquely truncated bladelets were foundthat are typical of the Kebaran. While the presence of several small triangles is notable, the frequency of
microliths among the tools is not as high as at Epi-Paleolithic sites on the coastal plain. This may signify thata different set of activities occurred at mountain sites than on the coastal plain. Further study is underway to
examine differentiation among layers II-IV and confirm whether these all of these set of layers can indeed be
attributed to the early Epi-Paleolithic. While only the top of layer V was uncovered and the lithic assemblage
from this unit is currently small, elements typical of the Levantine Aurignacian were noted, including
carinated tools and pieces bearing intrusive flat retouch from both lateral edges.
Paralleling our study of the new material we began technological analysis of the material from Ronen’s
excavation. We focused on Ronen’s Layer 8, which is considered to be one of a few “classic” Levantine
Aurignacian occupations (Goring-Morris and Belfer-Cohen 2006). Our technological analysis confirms
Aurignacian elements such as carinated scrapers/cores and pieces with Aurignacian retouch. We also foundextensive evidence for blade production characterized by the débitage frontal technique, shaping of ridge
blades and removal of overshots and core tablets. We hope to reconstruct the reduction sequences of blade
production characterizing the Levantine Aurignacian to answer such questions: To what extent does blade
production of the Levantine Aurignacian differ from the Ahmarian? (Davidzon and Goring-Morris 2003)
Was it constant in character throughout the three Levantine Aurignacian layers present at the site?
Complementing the lithic studies, a comparison of the marine shell assemblages from the Upper Paleolithic
and later periods illustrates certain trends. Upper Paleolithic inhabitants of Sefunim preferred small shells
such as Columbella rustica and scaphopods, while Neolithic inhabitants chose larger Glycymeris sp. andselected a more diverse array of mollusks (Table 2). Regardless of the period, shells were perforated at a
steady rate. Most Upper Paleolithic shells are small, suggesting that people brought them to the site forornaments and not food. Shell surface preservation is poor, indicating that specimens were collected frombeach deposits. The spectrum of marine shells allows us to infer that the marine ecosystem remained constantover time and is similar to today. As such, we interpret the variability in the assemblages of shells as anindication of different cultural perceptions (Bar-Yosef Mayer 2005).
Bone tools are few and mainly include fragments of perforators. While small pieces of ocher are commonly
found, one exceptionally large piece of red ocher was heavily abraded. With renewed excavation and further
studies underway, we expect to add significant new information about the transition from the Middle toUpper Paleolithic, as well as variability during the Upper Paleolithic.
References
Bar-Yosef Mayer DE (2005) The exploitation of shells as beads in the Palaeolithic and Neolithic of the Levant. Paléorient 31: 176–
185.
Davidzon A, Goring-Morris N (2003) Sealed in stone: The Upper Paleolithic early Ahmarian knapping method in the light ofrefitting studies at Nahal Nizzana XIII, Western Negev, Israel. Journal of Israel Prehistoric Society 33: 75–206.
Goring-Morris N, Belfer-Cohen A (2006) A hard look at the ‘Levantine Aurignacian’: How real is the taxon? In: Bar-Yosef O,Zilhão J (eds) Towards a definition of the Aurignacian. Instituto Português de Arqueologia, Lisboa, pp. 297–313.
Ronen A (1984) Sefunim prehistoric sites, Mount Carmel, Israel. BAR International Series 230, Oxford.
Stekelis M (1961) Iraq el-Barud, nouvelle grotte préhistorique au Mt. Carmel. Bulletin of the Research Council of Israel 10G: 302–
320.
Stiner MC, Kuhn SL, Güleç E (2013) Early Upper Paleolithic shell beads at Üçagızlı Cave I (Turkey): Technology and thesocioeconomic context of ornament life-histories. Journal of Human Evolution 64: 380–398.

041
The First RNMH Processes in West Asia

Fig. 1 Sefunim Cave. View of cave to the south with inset map
showing site location (yellow star) 10 km south of Haifa (Photo:
A. Fernandes).

Fig. 2 Sefunim Cave. Profile drawing showing stratigraphy of the 2013 excavation at the drip
line of the cave, as well as location of finds and micromorphological samples. Archaeologicalhorizon V corresponds to Ronen’s Levantine Aurignacian layer 8, but see Table 1 forcomplete correlation.
Table 1 Sefunim Cave. Cultural stratigraphy showing correlation of Ronen’s layers with thecurrent excavation.

PeriodModernNeolithic
Neolithic
Neolithic
Epipaleolithic
Epipaleolithic
Epipaleolithic/UP
Upper Paleolithic
Upper Paleolithic
Upper Paleolithic
sterileMP/UP
Middle Paleolithic

Industry

Ronen (1984)

1-4
5
6
7

Kebaran?
Kebaran?
Levantine AurignacianLevantine AurignacianLevantine Aurignaciansterile“transitional”
Levantine Mousterian

Current work

I

II
III
IV
V

8
9
10
11
12
13

Not yet excavated

Table 2 Sefunim Cave. Distribution of main types of marine shells and degree of perforationfrom both excavations using Ronen’s stratigraphy (Note: not all shells are presented here).
Columbella Nassarius Scaphopods Glycymeris Total Perforated Perforated
Periodrustica
gibbosulus
sp.
(n)
(%)
Surface and
1
1
2
3
33%
indeterminate
Neolithic and
12
4
13
31
39%
younger
UP Layer

14

042

8

1

1

MP/UP Layer
TOTAL SHELL

3

18

The First RNMH Processes in West Asia

5

10

14

29

12

41%

1

1

100%

64

26

41%

The Ahmarian in the context of the earlier Upper Palaeolithic
in the Near East
Nigel Goring-Morris1* and Anna Belfer-Cohen1

1. Institute of Archaeology, The Hebrew University of Jerusalem, Israel. *Email: nigel.goring-morris@mail.huji.ac.il

There is a general consensus that the Ahmarian technocomplex represents an endemic Upper Palaeolithic
entity that emerged in south-western Asia. Its entrenchment in the region is apparent over a long chronologicalspan and a wide geographic range, as is most especially apparent in the Levant. Notwithstanding diachronicand synchronic variability, its basic parameters have been widely recognized since it was first defined over 30
years ago. The defining characterization of the Ahmarian is based on certain intrinsic features as well as on
the absence of hallmarks of other Upper Palaeolithic entities in the region.
Our paper focuses on the techno-typological characteristics of the Ahmarian industries from an evolutionary
perspective, with specific examples from different phytogeographic settings within the Levant, in an attempt
to compare and contrast the variability present within the technocomplex and in relation to other Upper
Palaeolithic entities.

043
The First RNMH Processes in West Asia

The Upper Palaeolithic of Manot Cave, western Galilee, Israel

Ofer Marder1*, Israel Hershkovitz2 and Omry Barzilai3

1. Archaeology Division, Ben-Gurion University of the Negev, Israel; 2. The Dan David Laboratory, Sackler Faculty of Medicine, Tel Aviv
University, Israel; 3. Israel Antiquities Authority, Jerusalem, Israel. *Email: Mardero@bgu.ac.il

The Upper Palaeolithic of the Levant (45,000–22,000 BP) represents the full establishment of Modern
Human behavior in this region following the existence of both Modern Humans and Neanderthals
during the Middle Palaeolithic. The Levantine Upper Palaeolithic shares some similarities to its European
counterpart but otherwise is quite different. The Levantine Upper Palaeolithic is broadly divided into threechronological stages (Initial, Early, Late), with several cultural entities. While the Initial stage is the interface
between the Middle and the Upper Palaelithic, it is the Early Upper Palaeolithic when Modern Human
populations have fully established themselves in the region. The Early Upper Palaeolithic consists of two
cultural entities, “Ahmarian” and “Levantine Aurignacian”. The Ahmarian, dispersed throughout the Levant,is conceived as local tradition whereas the Aurignacian, constrained to few cave sites in the Mediterranean
woodland region, is considered an interference of European population/s.
In this paper we focuses on the chrono-cultural sequence at Manot Cave with special emphasis on the lithic
faunal as well as the human remains of the Upper Palaeolithic. Manot is a nearly-sealed, active karstic cave,located in a hilly landscape in the western Galilee, Israel. A recent excavation uncovered rich archaeologicalaccumulations attributed mainly to the Early Upper Palaeolithic period (both Ahmarian and Aurignacian
traditions) as well as a few Middle Palaeolithic, Initial Upper Palaeolithic and Epipalaeolithic remains.

044
The First RNMH Processes in West Asia

Living on the edge: The earliest Modern Human settlement of
the Armenian Highlands
Andreas Taller1*, Boris Gasparyan2 and Andrew W. Kandel3

1. Department of Early Prehistory and Quaternary Ecology, The Institute of Pre- and Protohistory and Mediaeval Archaeology, University
of Tübingen, Germany; 2. Institute of Archaeology and Ethnography, National Academy of Science, Armenia; 3. The Role of Culture in
Early Expansions of Humans (ROCEEH), Heidelberg Academy of Sciences and Humanities, University of Tübingen, Germany.
*Email: andreas.taller@uni-tuebingen.de

Aghitu-3 is the first stratified Upper Paleolithic cave site discovered in Armenia. The site is situated in the
Syunik volcanic region of the Southern Caucasus and has yielded three intact Upper Paleolithic horizons.
Each horizon contains archaeological finds including lithic artifacts, faunal remains and combustion
features. The site is also important because of its excellent preservation of paleoecological archives such as
micromammals, fish, birds, reptiles, amphibians, charcoal, pollen, non-pollen palynomorphs (NPPs) andtephras, which allow for a comprehensive interpretation of the climate and environment at the time the firstModern Humans populated the Armenian Highlands. Twelve geological horizons (GH) were identified, andcorrelate with seven archaeological layers (AH), as shown in Fig. 1. With dates ranging from 40-24,000 cal
BP, the site of Aghitu-3 offers us a glimpse into the settlement patterns of early Modern Humans during theearly and middle Upper Paleolithic of Armenia.
The layers that yielded the Upper Paleolithic material are AH VII, VI and III, while AH V and IV yieldedvery few artifacts. The distribution of lithic artifacts, faunal remains and charcoal is presented in Table 1. The
two lower find horizons AH VII and VI dated between about 40,000 and 33,000 cal BP, whereas the upper
find horizon, AH III, dates between 29,000 and 24,000 cal BP (Fig. 2).
This camp of hunters and gatherers very likely marks one of the earliest arrivals of anatomically Modern
Humans into the southern Caucasus. The lithic technology is aimed at producing laminar blanks following
a unidirectional logic, with bladelets predominating. The majority of these blanks are finely retouched along
one lateral edge, with semi-abrupt retouch also common. While true backed elements are present, they are in
the minority. Other retouched lithics such as scrapers, burins, splintered pieces and points are less frequent.
The lithic raw materials include obsidian and chert, both of high quality and from a variety of sources from as
far away as 250 km. The overall low number of artifacts, the low frequency of cores, and the comparably highnumber of retouched pieces suggest that this camp was used for short term stays rather than as a base camp.
Two awls and a broken eyed needle in AH III offer interesting perspectives on the use of bone tools andindicate that the inhabitants of Aghitu were making clothing, nets or bags. Most specimens of the faunalassemblage are mid-sized ungulates (wild sheep and wild goat) with equids also common. While evidence ofcarnivore activity is present in AH VII and VI, humans appear to have accumulated most of the fauna fromAH III. The genetic analysis of a complete canid skull and post-cranial elements suggests an affinity to wolf,
raising interesting questions about the timing of canid domestication in the Caucasus. Paleoenvironmental
indicators such as micromammal remains, pollen, charcoal and sedimentology suggest a cooler climate for
AH VII, but warm and humid conditions for AH VI. The climate became increasingly colder and drier withthe approach of the Last Glacial Maximum, as shown by the sequence from AH V to III.
Recent research at several Middle Paleolithic sites in Armenia has been successful in creating a newframework for understanding the rise and fall of Neanderthals in Armenia, for example, at the Middle
Paleolithic sites of Bagratashen-1, Hovk-1, Lusakert-1, Angeghakot-1, Kalavan-2 and Barozh-12. Thus the045

The First RNMH Processes in West Asia

well-stratified and recently excavated cave site of Aghitu-3 offers a unique perspective on the shift from
Middle to Upper Paleolithic technology with the arrival of anatomically Modern Humans in the Caucasus.
The 16,000 year time span represented by Aghitu-3 is particularly interesting as it allows us to investigate
diachronic as well as technological dimensions of the shift from the Middle to the Upper Paleolithic in theSouthern Caucasus. One of the many scientific challenges facing the research team will be the incorporationof the site into a wider regional context. Georgian sites such as Dzudzuana Cave and Ortvale Klde are ofgreat interest for comparison. Furthermore, the Armenian site of Kalavan-1 documents the late Upper
Paleolithic at the end of the last Pleniglacial (ca. 18-16,000 cal BP). Due to its unique stratification andage, Aghitu-3 will serve as the Armenian benchmark for the early and middle Upper Paleolithic. Finally,
the continuing analyses will make a crucial contribution to the development of a frame of reference for the
Caucasian Upper Paleolithic.
References
Kandel AW, Gasparyan B, Nahapetyan S, Taller A, Weissbrod L (in press) The Upper Paleolithic settlement of the Armenian
Highlands. In: Otte M (ed) Modes of contacts and displacements during the Eurasian Paleolithic, 29-31 May 2012, UISPP
Commission 8. ERAUL 140, Liege.

046

Fig. 1 Stratigraphy of Aghitu-3 (after Kandel et al. in press).

The First RNMH Processes in West Asia

Fig. 2 Calibrated AMS radiocarbon dates for bone andcharcoal from Aghitu-3.

Table 1 Distribution of lithic, faunal and charcoal finds in the archaeological horizons including data from the 2009-2013 fieldseasons analyzed through July, 2014.

Archaeological horizon (AH)
Area excavated (m2)
LITHICS
BlanksRetouched tools
Cores
Angular debris (chunks)
Small debitage (chips)
LITHIC subtotal
Retouch index (excluding chips)
FAUNA
Small mammals (SC1)
Small-medium mammals (SC2)Large-medium mammals (SC3)
Large mammals (SC4)
Fish
Hare
Fox
Wolf
FAUNA subtotal
CHARCOAL

III
40

IV
12

2408
564
66
128
1739
4905
17.8%

2
3
---5

12
491
422
16
15
4
1
1
962
821

--------5

V

VI
12

VII

6
4
---10

250
72
3
17
32
374
21.1%

1
4
1
1
---1
8
14

14
84
8
1
1
5
-2
115
227

12

TOTAL

%

94
9
5
4
-122
7.4%

2760
652
74
149
1771
5416

51%
12%
1.4%
2.8%
33%
100%

1
13
9
-24
1
--48
93

28
592
440
18
40
10
1
4
1133
1160

2.5%
52%
39%
1.6%
3.5%
0.9%
0.1%
0.4%
100%

4

047

The First RNMH Processes in West Asia

Typo-technological analysis of the Late Middle Paleolithic
bifacial industry of Qaleh Bozi Rockshelter, Central Iran
Fereidoun Biglari

Paleolithic Department, National Museum of Iran, Iran. Email: fbiglari@gmail.com

Introduction
Iranian Middle Paleolithic is mainly known from a number of excavated and surveyed cave and rocksheltersites in the Zagros Mountains which their lithic assemblages were the basis for defining of Zagros Mousterian.
An industry characterized by discoid and Levallois core methods of flake production. The Zagros Mousterian
assemblages characterized by intense core reduction and tool retouch that reflected in the small size of exhausted
cores and heavily retouched tools (Dibble 1984; Lindly 2005; Skinner 1965). As a result, the assemblages arerich in various types of side-scrapers such as single, double and convergent scrapers showing invasive and steep
retouch on their edges.
Recent research in a cluster of Middle Paleolithic cave and rockshelter sites near Isfahan, situated between Zagros
Mountains and central desert of Iran, produced lithic assemblages showing features that are new to IranianMiddle Paleolithic industries. This paper first briefly describes some results of excavations carried out in this site
complex and then it presents analysis of obtained lithic assemblages from Qaleh Bozi Rockshelter that provide
insights into the Middle Paleolithic lithic variability and human behavior in this transitional zone between highZagros Mountains and Iranian central deserts.
Qaleh Bozi sites
The Qaleh Bozi sites are situated about 25 km south-southwest of Esfahan, close to the inner foothills of theZagros range. The sites include a large rockshelter (QB 1), a large cave (QB 2) and a very small rockshelter (QB 3)
located between 1750 and 1810 meters above sea level, on the southern face of Qaleh Bozi Mountain.
Surface sampling and test excavations in QB2 and QB3 in 2005 and 2008 produced diverse vertebrate remains
and lithics artifacts, mostly in a fresh condition. (Biglari et al. 2009; Jaubert et al. 2010). Apart from small
excavated assemblages in 2005 and 2008 seasons, majority of the Middle Paleolithic material from Qaleh Bozi
2 found in a disturbed context. Test excavation in the Qaleh Bozi 3 yielded a small, but rich Middle Paleolithic
industry associated with scattered faunal remains.
Qaleh Bozi Rockshelter
Qaleh Bozi Rockshelter (QB 3) lies about 110 m to the south of the large cave of Qaleh Bozi (QB 2). The
shelter faces northwest, overlooking a steep rocky slope. It is a small rockshelter, 5 m deep, 3 m wide, with 8 m²
of interior floor space. Test units placed within a grid-coordinate system and six adjacent units (150×100cm)
were excavated to a depth of 180cm. The arbitrary excavation levels grouped into three major stratigraphic units,labeled I, II, and III from top to bottom.

048

In total, the excavated deposits yielded near 200 lithic artifacts and about 300 bone fragments and teeth whichcame from stratigraphic unit II. The majority of faunal assemblage belong to medium and large mammals. The

The First RNMH Processes in West Asia

only taxa clearly identified are small ruminants, with the presence of the teeth. The small ruminants can be thegazelle or wild caprini (sheep and goat). Despite high degree of bone fragmentation, presence of cut marks andpercussion marks on some of the specimens indicate involvement of human in their accumulation.
The lithic industry
Generally, the artifacts are in a very fresh condition with some specimen showing light patination. The mostabundant and locally available raw material is various types of cherts in pebble and cobble forms. These pebbles
and cobbles procured from the Zayandeh-Rud River, about 1500 m to the south and southeast of the sites. It
seems pebbles and cobbles were being brought to the site intact and without examination for quality.
The Qaleh Bozi 2 Rockshelter yielded a small collection of 196 artifacts. Large part of the assemblage (90.82 %)comes from excavation, and remaining pieces (9.18 %) found in backdirt pile from illegal excavation and fromprofile cleaning. The total weight of the assemblage is 925.5 grams. The lithic assemblage is largely composed ofdebris, chunks, shatters and flake fragments (60.21 %), followed by flakes and small flakes (19.25 %), tools (intact,broken and fragment) (15.32 %), and a small number of other categories such as pebble fragments. There are only
three cores and a core fragment (Table 1).
The assemblage is characterized by high frequencies of tools, flakes and flake fragments. There are several corefragments and a thick core-on flake. Bifacial reduction is an important technology in the industry, however truecore reduction is also used. There are few Levallois products in the assemblage. The tool group is heavily bifacial
and it is mainly include bifacial tools, Mousterian points and scrapers that accompanied by lower frequencies ofalternately retouched convergent tools, and other retouched pieces. The main characteristic of the tool group is
the high percentage of both bifacial and unifacial points which comprise about 40 percent of all retouched tools.
They are made of both flake blanks and pebbles. Typologically they can be assigned to Leaf shaped bifacial points,
unifacial point, partial bifacial, bifacial scrapers, and various scraper types: single side-scraper, double side-scraper,
convergent scraper-alternate retouch, Déjeté scraper and transverse scraper. There are also other tool types such as
Notched flake, borer, truncation, retouched pieces.
The lithic assemblage was examined for refits. The main aims were to examine the vertical integrity of the deposits
and to gain information about lithic technology and raw material use by Middle Paleolithic knappers during theirephemeral occupations in the site. As a result one conjoin group, one join group, and three associated groups wererecognized. Among the refits, the most informative is a refit series composed of five pieces. It includes four piecesof flakes and other categories, of which two directly reattached to a broken core. The specimens were separated
horizontally by less than 100 cm and vertically by less than 30 cm. This group suggests in situ knapping in the site.
A primary use-wear analysis was performed on three points from Middle Paleolithic layer at Institute ofPrehistory and Quaternary Geology, University of Bordeaux 1 (Claude et al. 2012). The three specimens
examined comprised two bifacial points and a Mousterian point from stratigraphic unit II. Use-wear traces were
found on two of the tools.
The first tool examined is a foliate point, showing oblique scars and meat polishes on the distal part of the leftedge which indicates its use in butchery. On the proximal part, especially on the central area, the ridges onboth faces show macro-rounding and polish, contrary to the distal part, where the ridges show no traces. These
traces may be the result of proximal hafting of the point. The second tool examined is a Mousterian point.
The left edge shows scarring on the lower face due to the cutting of soft to medium hard material. The scars
have a discontinuous distribution, are oblique, triangular and trapezoidal in morphology. Based on use-wear
characteristics, two points were used in butchery, most probably for cutting meat (Claude et al. 2012).

049

The First RNMH Processes in West Asia

Final remarks
Qaleh Bozi industry occupies a unique position in the Middle Paleolithic of Near East in general and Iran inparticular. The industry exhibit techno-typological characteristics which in some aspects resembling Zagros
Mousterian industries but also display certain features such as bifacial reduction which is not reported for IranianMiddle Paleolithic industries, nor from other Near Eastern Late Middle Paleolithic industries between Zagros
and eastern Mediterranean.
The question that the industry is an inclusion from other regions such as central Asia or Arabian Peninsula or
it is a local invention needs more new data from this region. The possibility of finding new sites with similarindustry cannot be ruled out on the basis of the present evidence. These new finds from Qaleh Bozi indicate thatthe region was not culturally uniform during Middle Paleolithic as supposed before (Lindly 2005; Skinner 1965).
References
Biglari F, Javeri M, Mashkour M, Yazdi Y, Shidrang S, Tengberg M, Taheri K (2009) Test excavations at the Middle Paleolithic
sites of Qaleh Bozi, southwest of Central Iran, a preliminary report. In: Otte M, Biglari F, Jaubert J (eds) Iran Palaeolithic. Proceedings of the XV World Congress UISPP, Lisbon. BAR International Series 1968, Archaeopress, Oxford, pp. 29-38.
Claud E, Biglari F, Jaubert J (2012) Preliminary use-wear analysis of several Middle Paleolithic points from Qaleh Bozi 3 rockshelter, Central Iran. Iranian Archaeology 3: 7-13.
Dibble HL, McPherron SP (2007) Truncated-faceted pieces: Hafting modification, retouch, or cores? In: McPherron SP (ed)Tool v. core: New approaches in the analysis of stone tool assemblages. Cambridge Scholars Publications, Cambridge.
Jaubert J, Biglari F, Crassard R, Mashkour M, Rendu W, Shidrang S (2010) Paléolithique moyen récent de la grotte de Qaleh
Bozi 2 (Esfahan, Iran): Premiers résultats de la campagne 2008. Iranian Archaeology 1(1): 21-31.
Lindly JM (2005) The Zagros Mousterian: A regional perspective. Anthropological Research Papers 56, Arizona State University, Tempe.
Skinner J (1965) The flake industry of southwest Asia: A typological study. PhD dissertation, Columbia University, New York/
University of Michigan, Ann Arbor.

Fig. 1 Bifacial tools and a point from stratigraphic unit II (arbitrary level 11 and 12), Qaleh Bozi.
Table 1 Qaleh Bozi 3 lithic assemblage composition.

050

Artifact category
Tools
FlakesFlakes (≤ 20mm)
Flake fragments
CoresCore fragment
Chunks and debris
Cobble/pebble fragments
Total

The First RNMH Processes in West Asia

Excavation Backdirt and profile
26
4
20
1
18
34
5
3
1
71
8
5
178
18

Total
30
21
18
39
3
1
79
5
196

%
15.32
10.7
9.18
19.9
1.54
0.5
40.31
2.55
100

The Middle to Upper Paleolithic transition in Zagros: The
appearance and evolution of Baradostian in the light of new
research
Sonia Shidrang

PACEA-PPP-Université Bordeaux 1, France. Email: sshidrang@gmail.com

The matter of Neanderthals disappearance and their replacement by Modern Humans or the transition from
Middle to Upper Paleolithic is an intensely debated topic in Paleolithic archaeology and paleoanthropologyof Europe and Western Asia. The emergence of Early Upper Paleolithic technologies signals significant
changes in human behaviors during Oxygen Isotope Stage 3 in Western Eurasia. Despite more than two
decades of intensive research, still no definitive agreement has been reached on the nature of this crucial
shift or its responsible factors (biological, environmental, socio-cultural or an intertwined process of multiple
factors). It is the beginning of the process through which anatomically Modern Humans dispersed widelyand its traces recorded in the variable lithic and bone industries reported from several regions in Europe
and Southwest Asia. For several decades, many of these lithic industries traditionally grouped under thetitle of Aurignacian technocomplexes which were considered as the marker of anatomically Modern Humandispersal. However, new lines of evidence and studies from involved geographical regions are essential to
improve such explanatory models and hypotheses. The Zagros Mountain range in the west of Iran with its
Intermountain eco-cultural niches is one of the areas that increasingly contribute to our knowledge of thetransition from Middle to Upper Paleolithic in Southwestern Asia. The major part of the information on
the Iranian Early Upper Paleolithic comes from the Zagros region or in other word from several cave androckshelter sites in the Intermountain valleys of Kermanshah and Khorramabad and a few sites in Fars
province in the southern Zagros (Fig. 1). The resemblance of Zagros Baradostian lithic industries with
Aurignacian technocomplexes of Europe and Levant and also the hypothesis that it evolved out of underlying
Zagros Mousterian engaged Baradostian as one of the potential candidates for ambiguous origin of
Aurignacian. On the basis of similarities observed in Warwasi assemblages and Aurignacian technocomplex,
Baradostian renamed “Zagros Aurignacian” and two main chronological phases described for it (Olszewski
and Dibble 1994, 2006). The earlier of these has been classiﬁed as the Early Zagros Aurignacian, and includes
a combination of Upper and Middle Paleolithic tools. Traditionally, the typo-technological continuities anddiscontinuities in lithic entities have been used to support the fundamental hypothesis of local evolution
versus acculturation in Paleolithic literature. In Zagros, Warwasi assemblage is presented as an example of atransitional industry, which presents a sequence of industrial development from Middle Paleolithic into the
Aurignacian. In addition to Warwasi, based on recent re-studies of lithic materials from 1965 excavation of
Yafteh cave, the existence of two main typo-technological phases were described in the Yafteh Baradostiansequence (Bordes and Shidrang 2009) which share certain similarities with two phases of Warwasi but also
have differences that all are described in this paper (Figs. 2 and 3). A series of new radiocarbon dates suggeststhat the Baradostian in Yafteh attributes to an age between 24,500 and 36,000 14C BP (Table 1) and a more
recent results indicate a single chronological signal of approximately 33,500 14 C BP for the major part of
Yafteh sequence (Otte et al. 2007, 2011). Radiocarbon estimations from main occupation of Yafteh appear topre-date the chronological range of considerable Levantine Aurignacian sequences and overlap some of the
Early Ahmarian ones (Otte et al. 2011).
This paper examines the lithic-based dominant hypothesis of continuity in Zagros through a more technology
oriented view and put together all the evidence to build a broader overview of the Early Upper Paleolithic in
Zagros and its industrial evolution. It also looks into the other lines of evidence of technological innovation

051

The First RNMH Processes in West Asia

and new behavioral patterns such as relatively high exploitation of organic materials alongside lithic
manufacturing strategies in Baradostian (Zagros Aurignacian) compared to Mousterian and emphasizes
on different perspectives of change (Fig. 4). The main objective of the paper is the nature and extent of
behavioral change in the beginning of the Early Upper Paleolithic in Zagros and implications for significant
increase of behavioral complexity. For this purpose a critical review of existing hypotheses of Zagros Middle
to Upper Paleolithic transition is provided and new data from a current technological study of rich and wellpreserved Yafteh lithic assemblages opens new perspectives on the subject. Based on this recent detailed
analysis, which included both 2005-2008 assemblages of Yafteh cave test excavation, stratigraphical andother archaeological remains, there are three distinguishable cultural phase in Yafteh sequence. Also based on
taphonomical analysis, we now have evidence to suggest that most parts of the Yafteh deposits and its units
are well-preserved and contain reliable cultural information which contributes to a better understandingof techno-economical changes throughout the sequence. As the sequence of Yafteh leads us, we can trace
the evolution of the Baradostian industries throughout its core management toward a more volumetricshape and more complex and diverse reduction sequences. Thus, incorporating the new information resulted
from a current typo-technological and taphonomical study of the Yafteh cave assemblages into the existingknowledge provides us with a better reconstruction of the Baradostain sequence in the Zagros.
References
Bordes JG, Shidrang S (2009) La sequence baradostienne de Yafteh (Khorammabad, Lorestan, Iran). In: Otte M, Biglari F, Jaubert J(eds) Le Paleolithique d’Iran. Archaeopress, Oxford, pp. 85-100.
Olszewski DI, Dibble HL (1994) The Zagros Aurignacian. Current Anthropology 35: 68-75.
Olszewski DI, Dibble HL (2006) To be or not to be Aurignacian: The Zagros upper Paleolithic. In: Bar-Yosef O, Zilhao J (eds)
Towards a deﬁnition of the Aurignacian. Trabalhos de Arqueologia 45, American School of Prehistoric Research/Instituto Portugues
de Arqueologia, Lisbon, pp. 355-373.
Otte M, Biglari F, Flas D, Shidrang S, Zwyns N, Mashkour M, Naderi R, Mohaseb A, Hashemi N, Darvish J, Radu V (2007) The
Aurignacian in the Zagros region: New research at Yafteh cave, Lorestan, Iran. Antiquity 81: 82-96.
Otte M, Shidrang S, Zwyns N, Flas D (2011) New radiocarbon dates for the Zagros Aurignacian from Yafteh cave, Iran. Journal of
Human Evolution 61: 340-346.

052

Fig. 1 The location of main known Early Upper Paleolithic
sites of Iran (Basemap courtesy of NASA’s Visible Earth, http://
visibleearth.nasa.gov/).

The First RNMH Processes in West Asia

Fig. 2 Baradostian bladelets (Type B); 2. Baradostianbladelets (Type A); 3. Arjeneh points (Drawings: S.
Shidrang).

Fig. 4 The earliest known evidence of symbolic and relatively
complex behaviors in Early Upper Paleolithic of Iran (Photos:
Shidrang and Biglari).

Fig. 3 The main part of bifacial bladelet cores reduction
sequence in Yafteh cave assemblages (Drawings: S.
Shidrang).

Table 1 Table summarizing all 14 C dating and calibrated results from Yafteh cave (Otte et al. 2011).

053
The First RNMH Processes in West Asia

DAY 3: December 2, 2014

Environmental Backgrounds for RNMHTiming of the RNMH in Europe
Global Processes of RNMH

Presentation of the PaleoGeo database with Google Earth

Wonsuh Song1*, Yasuhisa Kondo2 and Takashi Oguchi1

1. CSIS, The University of Tokyo, Japan; 2. Research Institute for Humanity and Nature, Japan. *Email: wonsuhsong@gmail.com

Paleoenvironmental studies cover various fields such as paleohydrology, geomorphology, paleooceanology,
paleobiology, paleoclimatology, and chronology. It is difficult for an individual researcher to collect andcompile enormous data regarding these fields. We have been compiling portal data and presenting them using
a web-based geographical information system (Web-GIS) called PaleoGeo for the multidisciplinary project“Replacement of Neanderthals by Modern Humans”.
The PHEIMS (Paleo-Hydrology and Environment Internet Map Server), a precursor of the PaleoGeo, wasdeveloped in the late 1990s by one of the co-authors (T. O.), using ESRI ArcView IMS as a main server software
package. It contained information from papers in international journals of earth and Quarternary sciences
published during the mid-1990s to 2002. The PaleoGeo is being developed to revive and update the PHEIMS.
We improved the user interface and data quantity. The new system uses ESRI ArcGIS Server 10 and to reduce
redundancy of the data, a relational database management system (RDBMS) is applied. Fig. 1 shows the interfaceof the PaleoGeo. Search results are shown on the map with red pins. These visualized metadata are indispensible
for many researchers, for example, the Ostracod Metadatabase of Environmental and Geographical Attributes(Horne et al. 2011), and the Meta-database for Recent Paleolimnological Studies (Battarbee et al. 2007).
Google Earth is one of the most powerful tools providing high-resolution aerial imagery for the entire world(Visser et al. 2014; Demirci et al. 2013). Google Earth is not considered as a true GIS due to its limited spatial
analytical operations (Patterson 2007).However, it has been used by many professionals and individuals since it
was released in 2005 (Google 2007).
We applied this relatively recent tool for our research. First, all the collected data have to be converted to a
Keyhole Markup Language (KML) file in the ArcGIS. KML is an XML-based language for defining thegraphic display of spatial data in applications such as Google Earth and ArcGIS Explorer (Conroy et al. 2008;
ESRI 2012). The extension of the file should be “.kmz”. Second, if a user opens Google Earth and drags the fileto the “Places”, then the data will be shown. Fig. 2 shows the PaleoGeo database with Google Earth. Only one
click on the display enables showing all the details including the journal name, information about each paper(authors, title, volume, year, and page numbers), theme, subtheme, keywords, DOI (Digital Object Identifier), andperiod (era). We found some advantages using Google Earth: 1) easier interface, 2) easier access, and 3) low cost.
It is user-friendly and does not need a long-term training or software fee. Bird’s eye view and street view are alsoavailable in the same operation tool (Fig. 3). Google Earth provides search function in its user interface; however,it does not search the whole of the uploaded data but only addresses.

056

The aim of the project behind the database is to reconstruct the distribution of Neanderthals and Modern
Humans in time and space in relation to past climate change. We have been collecting information fromalmost three thousand articles of 13 journals regarding paleoenvironmental research (i.e., Boreas, Catena,
Climatic Change, Earth Surface Processes and Landforms, Geomorphology, Journal of Quaternary Science,
Palaeogeography, Palaeoclimatology, and Palaeoecology, Quaternary International, Quaternary Research,
Quaternary Science Reviews, The Holocene, and The Journal of Geology). The topics of the articles were

Environmental Backgrounds for RNMH

classified into six themes (paleohydrology, earth surface processes and materials, paleooceanology, paleobiology,
palaeoclimatology, and chronology) and 19 subthemes (hydrology, flood, fluvial, glacier, fluvial/glacier,sedimentology, soil, slope process, periglacial, peat land, eolian, sea-level, biology, vegetation, zoology, vegetation/
zoology, archaeology, climate, atmosphere, and chronology). PaleoGeo still has a very useful function, such as
quick search function, in the user interface. However, it is less user-friendly comparing to Google Earth. In
addition, the number of the paleoenvironmental studies is growing rapidly and we have to effectively cover themas many as possible. We plan to simplify the input data (latitude, longitude, title and DOI only) to include more
publications. So far information about >7600 sites has been collected and the number is increasing. The collected
data are accessible via the internet (http://neangis.csis.u-tokyo.ac.jp/paleogeo/). The KMZ file of the data which
can be used with Google Earth will be open in a near future. Google Earth can be a better tool for sharing themetadata.
References
Battarbee RW, Morley D, Bennion H, Simpson GL (2007) A meta-database for recent paleolimnological studies. PAGES News 15:
23-24.
Conroy GC, Anemone RL, Regenmorter JV, Addison A (2008) Google Earth, GIS, and the Great divide: A new and simple method
for sharing paleontological data. Journal of Human Evolution 55: 751-755.
Demirci A, Karaburun A, Kilar H (2013) Using Google Earth as an educational tool in secondary school geography lessons.
International Research in Geographical and Environmental Education 22(4): 277-290.
ESRI (2012) An overview of the To KML toolset: ArcGIS Resources, ArcGIS Help 10.1.
Google (2007) Introducing Google Earth outreach Google, Inc. Press Release. Retrieved fromhttp://googlepress.blogspot.jp/2007/06/introducing-google-earth-outreach_26.html
Horne DJ, Curry BB, Delorme LD, Martens K, Smith AJ, Smith RJ (2011) OMEGA: The Ostracod Metadatabase ofEnvironmental and Geographical Attributes. Joannea Geologie und Paläontologie 11: 80-84.
Patterson TC (2007) Google Earth as a (not just) geography education tool. Journal of Geography 106(4): 145-152.
Visser V, Langdon B, Pauchard A, Richardson DM (2014) Unlocking the potential of Google Earth as a tool in invation science.
Biological Invations 16: 513-534.

Fig. 1 Interface of the PaleoGeo.

057
Environmental Backgrounds for RNMH

Fig. 2 Presentation of the PaleoGeo database with Google Earth.

Fig. 3 Bird`s eye view and street view available on Google Earth.

058
Environmental Backgrounds for RNMH

Regional last glacial climate synthesis for the Eastern
Mediterranean
Stephen P. Obrochta1*, Yusuke Yokoyama1 and Hodaka Kawahata1
1. Atmosphere and Ocean Research Institute, The University of Tokyo. *Email: obrochta@aori.u-tokyo.ac.jp

We present a regional climate synthesis for the Eastern Mediterranean region based on a variety of records
including lake sediment core pollen and hydrologic indexes, as well as speleothem stable isotopes. We both
examine spatial trends and gradients by considering all records individually and create a stacked record whichis compared to the North GRIP ice core.
Before stacking, care was taken to ensure that the age models of each record were consistent. For older
records that used the GISP2 the Meese-Sowers 1994 age scale (Meese et al. 1997), we updated their age
models to the GICC05 model-extended chronology (Andersen et al. 2006; Rasmussen et al. 2006; Svensson
et al. 2008; Wolff et al. 2010) using our recently developed function (Obrochta et al. 2014).
Results show in general good agreement with Greenland and North Atlantic climate, with generally
variations corresponding to Dansgaard-Oeschger oscillations. Cool and dry conditions are common during
stadials, and warm and moist conditions are observed during interstadials.
References
Svensson A, Andersen KK, Bigler M, Clausen HB, Dahl-Jensen D, Davies SM, Johnsen SJ, Muscheler R, Parrenin F, Rasmussen SO,
Röthlisberger R, Seierstad I, Steffensen JP, Vinther BM (2008) A 60 000 year Greenland stratigraphic ice core chronology. Climateof the Past 4(1): 47–57.
Andersen KK, Svensson A, Johnsen SJ, Rasmussen SO, Bigler M, Röthlisberger R, Ruth U, Siggaard-Andersen, M-L, Peder
Steffensen J, Dahl-Jensen D, Vinther BM, Clausen HB (2006) The Greenland ice core chronology 2005, 15-42 ka. Part 1:
Constructing the time scale. Quaternary Science Reviews 25(23-24): 3246–3257.
Meese DA, Gow AJ, Alley RB, Zielinski GA, Grootes PM, Ram M, Taylor KC, Mayewski PA, Bolzan JF (1997) The Greenland Ice
Sheet Project 2 depth-age scale: Methods and results. Journal of Geophysical Research 102(C12): 26411–26423.
Obrochta SP, Yokoyama Y, Morén J, Crowley TJ (2014) Conversion of GISP2-based sediment core age models to the GICC05
extended chronology. Quaternary Geochronology 20(0): 1–7.
Rasmussen, SO, Andersen KK, Svensson AM, Steffensen JP, Vinther, BM, Clausen, HB, Siggaard-Andersen M-L, Johnsen SJ,
Larsen LB, Dahl-Jensen D, Bigler M, Röthlisberger R, Fischer H, Goto-Azuma K, Hansson ME, Ruth U (2006) A new Greenland
ice core chronology for the last glacial termination. Journal of Geophysical Research: Atmospheres 111(D6).
Wolff EW, Chappellaz J, Blunier T, Rasmussen SO, Svensson A (2010) Millennial-scale variability during the last glacial: The icecore record. Quaternary Science Reviews 29(21-22): 2828–2838.

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Vegetation change in Europe during the Late Pleistocene
dispersal of Modern Humans
Ryouta O’ishi1,2*, Wing-Le Chan2 and Ayako Abe-Ouchi2,3

1. National Institute of Polar Research, Japan; 2. Atmosphere and Ocean Research Institute, The University of Tokyo, Japan; 3. Japan
Agency of Marine-Earth Science and Technology, Japan. *Email: ryo@aori.u-tokyo.ac.jp

The spatial and temporal variation of vegetation distribution is an important factor in understanding the
migration of Modern Humans during the Late Pleistocene. Geological evidence such as proxy data is spatiallylimited and it is difficult to reconstruct vegetation distribution. In order to do so, it is more effective to not only
make use of the geological evidence but also to combine it with numerical simulations. Team B02 is trying to
reproduce stadial-interstadial climate change, typically known as the Dansgaard-Oeschger events, by a GCM(General Circulation Model).
In the present study, we developed a new method to reproduce past vegetation distribution by applying past
climate information from the GCM to a DGVM (Dynamical Global Vegetation Model) with a practicalresolution which is comparable to scales associated with the anthropological evidence. We will show preliminary
results for vegetation changes which correspond to differences between a typical interstadial and typicalstadial climate. We also discuss the reliability of the obtained vegetation distribution by comparing with thepaleovegetation reconstruction based on sediment cores. We also discuss the advantages and limitations of thismodelling method.
References
Sitch S, Smith B, Prentice IC, Arneth A, Bondeau A, Cramer W, Kaplan JO, Levis S, Lucht W, Sykes MT, Thonicke K, Venevsky S(2003) Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the lpj dynamic global vegetation model.
Global Change Biology 9(2): 161–185.

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Cost surface analysis based on ecological niche probability
to estimate relative rapidity of the dispersals of early Modern
HumansYasuhisa Kondo

Research Institute for Humanity and Nature, Japan. Email: kondo@chikyu.ac.jp

Introduction
Timing and routes of the dispersal of early Modern Humans to Eurasia have actively been discussed (Petraglia etal. 2010; Mellars and French 2011; Dennel and Petraglia 2012; Bar-Yosef and Belfer-Cohen 2013; Boivin et al.
2013; Digandžic and McPherron 2013). In order to better understand the timing and process of migration fromthe archaeological and ecological viewpoints, the Replacement of Neanderthal by Modern Humans (RNMH)
project is carrying out an interdisciplinary research involving archaeologists, geochronologists, climatologists,
geomorphologists, and computer scientists (Kondo et al. 2013). The research group has developed a cyclopedic
database on archaeological sites and lithic industries in Africa, Eurasia, and Oceania in 200 to 20 kya (Kondo etal. 2012a), a technique of the statistical refinement of radiocarbon dates (Omori and Sano this volume), and a
new palaeoclimate model (Abe-Ouchi et al. 2013). These progresses made it possible to simulate the ecologicalniches of the human species through an ecological niche model (ENM), or a machine-learning model using
the location of sites and environmental indices (such as surface air temperature, precipitation, and elevation) as
heuristic operator (Kondo et al. 2012b). Furthermore, the ecological niche model can contribute to the estimationof the dispersal rate and optimal migration routes by means of the cost surface analysis in the following way.
Cost surface analysis
Cost surface analysis is a computer-based method to find a route that minimises the total cost of moving between
two locations on an accumulated cost surface (Conolly and Lake 2006: 294). Such a route is called least-cost
path. An accumulated (or cumulative) cost surface is created by a friction model, in which a friction value, or an
index of moving cost, is allocated to each spatial pixel. In the conventional cost surface models (Gorenflo andGale 1990; Tobler 1993; Van Leusen 2002; Neteler and Mitasova 2008: 381; Kondo and Seino 2010; Kondo
et al. 2011), slope of terrain is usually used as friction value. At this point, we can use the inverse of the nicheprobability as alternative friction.
Ecological niche model
The niche probability of past human species can be calculated by ENM, under the assumption that humanbehavior was, particularly in prehistory, largely influenced by the environmental factors (Banks et al. 2008a,
2008b, 2013). In practice, the niche probability is calculated for each spatial pixel, based on (1) the location of
known archaeological sites as occurrence data and (2) environmental factors such palaeoclimate (temperatureand precipitation), palaeovegetation (biome) and palaeotopography (elevation, slope and aspect). Some ENM
software packages, including MaxEnt (maximum entropy model), output the percentage of contribution foreach environmental factor, and therefore it is possible to identify and evaluate environmental constraints to thegeographic expansion of human populations. Based on this thought, the authors applied ENM to Palaeolithic
stone tool industries in Europe and Siberia at 50–46 kya, the time period during which the first Modern
Humans are presumed to have appeared in those regions, under the assumption that stone tool groups may reflect
different human groups in terms of subsistence strategy. The preliminary results suggested that the population

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using Emiran and related industries (Bohunician and Bachokirian) were likely to construct their niches in thegeographic zones where the long-term variability of the coldest month temperature was larger than in those
occupied by the population using Late Mousterian, Szeletian and Châtelperronean stone tool industries.
The study area (20°W to 160°E, 0° to 90°N) covered the most parts of Eurasia. Archaeological sites wereclassified into five lithic industry groups––(1) the Late Mousterian in the Iberian Peninsula at that period,and (2) the Châtelperronean in Western Europe, (3) the Szeletian in central Europe, (4) the Uluzzian in the
Italian Peninsula, and (5) Emiran-related industries, including the Emiran in the Levant, the Bachokirian in the
Balkans, the Bocunician in Central and Eastern Europe, and the Kara Bom culture in the Altai.
Regarding the palaeoenviromental data, ETOPO-1 was used for digital elevation model. Temperature andprecipitation at 50–46 kya were temporarily approximated by the 6 kya climate model of MIROC 3.2 because
they were under calculation. Palaeoclimatic variability was indexed for each topographic cell by the difference
between values at 21 kya (or Last Glacial Maximum) and 0 kya (present days) of the MIROC 3.2.2.
The maximum entropy model (MaxEnt; Phillips et al. 2006) was applied for calculating the niche probability. All
models show that the diachronic variability in the coldest month temperature most contributes to the model. The
geometric mean of the niche probability values of two different industries were considered as the niche overlap
rate. The results show that high niche overlap rates are observed in Central Europe, where the Szeletian and the
Bohunician were present. If we assume that the Kara Bom industry in the Altai derived from the Emiran in theLevant, the model indicated three possible niche corridors through Central Asia.

062

References
Abe-Ouchi A, Saito F, Kawamura K, Raymo ME, Okuno J, Takahashi K (2013) Insolation-driven 100,000-year glacial cycles andhysteresis of ice-sheet volume. Nature 500: 190-193.
Banks WE, d’Errico F, Peterson AT, Vanhaeren M, Kageyama M, Sepulchre P, Ramstein G, Jost A, Lunt D (2008) Human ecological niches and ranges during the LGM in Europe derived from an application of eco-cultural niche modeling. Journal of Archaeological Science 35: 481-491.
Banks WE, d’Errico F, Peterson AT, Kageyama M, Sima A, Sánchez-Goñi MF (2008) Neanderthal extinction by competitive exclusion. PLoS ONE 3: e3972. doi: 10.1371/journal.pone.0003972.
Banks WE, d’Errico F, Zilhão J (2013) Human-climate interaction during the Early Upper Paleolithic: Resting the hypothesis of an
adaptive shift between the Proto-Aurignacian and the Early Aurignacian. Journal of Human Evolution 64: 39-55.
Bar-Yosef O, Belfer-Cohen A (2013) Following Pleistocene road signs of human dispersals across Eurasia. Quarternary International
285: 30-43.
Boivin N, Fuller DQ, Dennell R, Allaby R, Petraglia MD (2013) Human dispersal across diverse environments of Asia during theUpper Pleistocene. Quaternary International 300: 32-47.
Conolly J, Lake M (2006) Geographical information systems in archaeology. Cambridge UP, Cambridge.
Dennel R, Petraglia MD (2012) The dispersal of Homo sapiens across southern Asia: How early, how often, how complex? Quarternary Science Review 47: 15-22.
Digandžic T, McPherron SP (2013) Demography and the demise of Neandertals: A comment on ‘Tenfold population increase inWestern Europe at the Neandertal-to-Modern Human transition’. Journal of Human Evolution 64: 311-313.
Gorenflo LJ, Gale N (1990) Mapping regional settlement in information space. Journal of Anthropological Archaeology 9: 240-273.
Kondo Y, Ako T, Heshiki I, Matsumoto G, Seino Y, Takeda Y, Yamaguchi H (2011) FIELDWALK@KOZU: A preliminary reportof the GPS/GIS-aided walking experiments for remodeling prehistoric pathways at Kozushima Island (East Japan). In: Jerem E
et al (eds) On the road to reconstructing the past: Computer applications and quantitative methods in archaeology. Proceedings ofthe 36th international conference. Archaeolingua, Budapest. http://proceedings.caaconference.org/files/2008/CD44_Kondo_et_al_
CAA2008.pdf. Accessed 23 July 2014.
Kondo Y, Kadowaki S, Kato H, Naganuma M, Ono A, Sano K, Nishiaki Y (2012a) Network computing for archaeology: A casestudy from the "Replacement of Neanderthals by Modern Humans” lithic industry database project. In: Zhou M et al (eds) Revive
the past: Proceeding of the 39th conference on computer applications and quantitative methods in archaeology, Beijing, 12-16 April
2011. Amsterdam UP, Amsterdam.
Kondo Y, Omori T, Verhagen P (2012b) Developing predictive models for palaeoanthropological research: A preliminary discussion.

Environmental Backgrounds for RNMHTechnical Report, Department of Computer Science, Tokyo Institute of Technology, TR12-0001.
Kondo Y, Sano K, Kadowaki S, Naganuma M, Omori T, Yoneda M, Nishiaki Y (2013) Eco-cultural niche modelling for the replacement of Neanderthals by Modern Humans based on a collaboration of the archaeology and palaeoenvironment teams. In: Terashima
H (ed) The 8th conference on replacement of Neanderthals by Modern Humans: Testing evolutionary models of learning.
Kondo Y, Seino Y (2010) GPS/GIS-aided walking experiments and data-driven travel cost modeling on the historical road of
Nakasendō-Kisoji (Central Highland Japan). In: Frischer B et al (eds) Making history interactive: Computer applications and quantitative methods in archaeology (CAA). Proceedings of the 37th international conference, Williamsburg, Virginia, United States ofAmerica, March 22-26, 2009. Archaeopress, Oxford. http://proceedings.caaconference.org/files/2009/21_Kondo_Seino_CAA2009.
pdf. Accessed 25 July 2014.
Mellars P, French, JC (2011) Tenfold population increase in Western Europe at the Neandertal–to–Modern Human transition. Science 333: 623-627.
Neteler M, Mitasova H (2008) Open source GIS: A GRASS GIS Approach. 3rd edn. Springer, London.
Petraglia MD, Haslam M, Fuller DQ, Boivin N, Clarkson C (2010) Out of Africa: New hypotheses and evidence for the dispersal of
Homo sapiens along the Indian Ocean rim. Annals of Human Biology 37: 288-311.
Phillips SJ, Anderson RP, Schapire RE (2006) Maximum entropy modeling of species geographic distributions. Ecological Modelling190: 231-259.
Tobler W (1993) Three presentations on geographical analysis and modeling. Technical Report 93/1. http://www.ncgia.ucsb.edu/
Publications/Tech_Reports/93/93-1.PDF. Accessed 25 July 2014.
Van Leusen PM (2002) Pattern to process: Methodological investigations into the formation and interpretation of spatial patterns inarchaeological landscapes. PhD dissertation, University of Groningen.

Fig. 1 Niche probability for the ProtoAurignacian lithic industry group at 43/42 to
40/39 kya, in response to the cooler climate
inferred from the hosing experiment.

Fig. 2 Relative accumulated cost for theProto-Aurignacian lithic industry group at
43/42 to 40/39 kya, in response to the cooler
climate inferred from the hosing experiment.

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Introduction to the “PalaeoChron” Project: Chronologicalmapping of the Middle and early Upper Palaeolithic of Eurasia
Katerina Douka1* and Tom Higham1

1. Research Laboratory for Archaeology and the History of Art (RLAHA), School of Archaeology, University of Oxford, UK.
*Email: katerina.douka@rlaha.ox.ac.uk

Recent advances in chronometric dating
A reliable chronology is crucial if we are to understand aspects of human evolutions, such as the spread ofModern Human populations from one or more core areas in Africa, the extinction of archaic human populations
living in Eurasia and the time of overlap between various human groups.
The last decade has seen major advances in the application of radiocarbon and other dating methodologies
focusing on “old” samples (prior to 30 ka) and therefore directly affecting the construction of Palaeolithic
chronologies. These advances include the development of much better protocols for purifying samples of bone
collagen using ultrafiltration, of charcoal using ABOx-SC and of shell using the CarDS method, prior to AMS
14C measurement.
Additionally, an internationally-agreed calibration curve extending to 50 ka for the conversion of radiocarbon
ages to calendar ones, as well as the wider use of Bayesian statistics, are crucial advances towards high-resolution
chronologies. Developments of the same scale and significance affect the application of luminescence dating on
sediment and heated flint.
The “PalaeoChron” Project (2013-2018)
In 2012, our group was awarded a major European grant,“PalaeoChron”- Precision dating of the Palaeolithic, tocontinue and expand previous work on dating late Middle and early Upper Palaeolithic sequences across Eurasia.
This work, which started in 2003 focusing originally on the British Palaeolithic, expanded between 2006-2011 toinclude a large number of Palaeolithic sites in west and southern Europe and the Near East.
With “PalaeoChron” we plan to expand our study area further to the east and into greater Eurasia to include sites
located in Eastern Europe, Russia and Crimea, Central Asia and the Levant, the Altai and Transbaikal of Siberia,
and Southeast Asia. We hope that with the wider participation of archaeologists from other regions, including East
Asia, we will be able to construct a broader chronological framework and a spatio-temporal model for placing inmajor cultural and biological changes observed in the archaeological record over the last 100 ka.
In this talk, we will briefly discuss the aims of PalaeoChron and its sub-projects (radiocarbon dating, luminescencedating, single amino-acid dating, ZooMS) and we will present some preliminary results relevant to the replacementof Neanderthals by Modern Humans.

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Timing of the RNMH in Europe

Radiocarbon dating the final Mousterian of Eurasia using
advanced chemistry methods
Tom Higham

Oxford Radiocarbon Accelerator Unit, RLAHA, University of Oxford, UK. Email: Thomas.higham@rlaha.ox.ac.uk

To understand the disappearance of the Neanderthals, a reliable chronology is required. This has largely
eluded science thus far; despite many radiocarbon determinations being obtained from sites dating to theMiddle to Upper Palaeolithic transition, our knowledge of the timing of this is opaque. The western European
archaeological record constitutes the best evidence there is for the prehistoric replacement of one species ofhuman by another, so deciphering the signals of this transition assumes importance in understanding what
happened. For example, exploring the dates of the disappearance of Neanderthals and the arrival of Modern
Humans might help us to understand the possibilities for overlap between the two groups, and therefore
whether there was any genetic or genetic exchange between the two populations.
AMS dating material dating older than 30,000 BP is challenging, but work undertaken in Oxford over thelast decade has improved the dating of material from this period. We have developed aspects of our pretreatment chemistry, particularly the purification of bone collagen using ultrafiltration and in the dating of
single amino acids from bone, as well quantifying our background limits and corrections.
Within a large project funded by the NERC in the UK, we have been dating over 400 samples of bone,shell and charcoal from more than 50 key Palaeolithic sites in 10 countries. The main focus has been onsites with a succession of contexts containing lithic industries attributed to the Mousterian, Châtelperronian(both seemingly associated with Neanderthals), Uluzzian (now thought to be anatomically Modern Humans(AMH)), Aurignacian and Gravettian (associated with AMHs).
In this paper we will discuss the emerging chronology for the disappearance of the Neanderthals bypresenting results from some of the key sites in France, Germany, Italy, Spain and Belgium. We use Bayesianmodelling to derive posterior density estimates for the final disappearance data of Neanderthals in eachgeographic location we have studied and consider the results in a spatio-temporal model. We are now
working on a range of sites dating to this period from sites further afield in Eurasia. We will describe also
some of the latest results from this work.

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Timing of the RNMH in Europe

Spatiotemporal pattern of the Middle to Upper Paleolithic
transitional lithic industries in Europe: A new approach forreliability assessment of radiocarbon data
Takayuki Omori1* and Katsuhiro Sano1

1. The University Museum, The University of Tokyo, Japan. *Email: omori@um.u-tokyo.ac.jp

Discussion based on a spatiotemporal distribution of archaeological sites requires extensive radiometric data in a database.
However, it is crucial to pay attention to a lot of problematic or unreliable data compiled in the database. We developeda systematic data assessment process for the database to reveal the probable spatiotemporal pattern of lithic industries
related to the Neanderthal and Modern Humans. In this presentation, we will present the new data assessment process
and the spatiotemporal distribution of transitional lithic industries in Europe using the data mining of the RNMH
database (Nishiaki 2012).
Recently, the accuracy in 14C dating especially against Paleolithic samples is increased by the development of thesample preparation procedures, such as ultrafiltration for bone (Bronk Ramsey et al. 2004) ABOx-stepped combustion
for charcoal (Bird et al. 1999) and carbonate density separation for shell (Douka et al. 2010). With the advent of the
improved preparation methods, the absolute chronologies of key archaeological sites around the world were revised
rapidly. In the new data assessment, we focused on the age differences between the new and conventional pretreatments.
Assuming that the 14C age from improved pretreatments has higher reliability than the 14C age from the conventional
pretreatments, the reinterpretation of the low-reliability 14C age and its error is performed. To construct the dataassessment process, we collected sets of “new” 14C age and “conventional” 14C age from the same sample from technicalpapers comparing preparation methods, and investigated the age difference and variation between pretreatments. As aresult, the systematical tendencies was revealed in ultrafiltration and ABOx-SC, and then represented as data assessment
formulas. By using the formulas, we can obtain the 14C age presumed to have been dated by the improved pretreatmentsfrom “conventional” 14C age. Although the error margin becomes larger than the prior, it is expected that the reliability of
14
C age has increased. To evaluate the assessment process, the chronological sequences of Geißenklösterle, Fumane Caveand Campanian Ignimbrite, which had been constructed by problematic “conventional” 14C ages, were re-examined bythis approach. The results shown good agreements with the recent revised data, and suggested the benefit of the process.
Applying this approach to the RNMH database, we revised the chronology of the transitional lithic industries in Europe.
Our examination focused on 9 lithic industries to discuss the process of extinction of Neanderthals and expansion ofModern Humans: the final Mousterian, Szeletian, Bachokirian, Bohunician, Uluzzian, Chatelperronian, Jerzmanowician,Proto-Aurignacian and Early Aurignacian. Based on all of the data in the RNMH database, we will illustrate thespatiotemporal pattern of the traditional industries, and show the new perspective about the replacement of Neanderthal
by Modern Humans.

066

References
Nishiaki Y (2012) The archaeological database for the RNMH Project. In: Akazawa T, Nishiaki Y (eds) RNMH 2012 -The firstinternational conference on the replacement of Neanderthals by Modern Humans: Testing evolutionary models of learning. RHMH
Project Group, Tokyo, pp. 92-93.
Bronk Ramsey C, Higham TFG, Bowles A, Hedges REM (2004) Improvements to the pretreatment of bone at Oxford. Radiocarbon
46: 155-163.
Bird MI, Ayliffe LK, Fifield LK, Turney CM (1999) Radiocarbon dating of “old” charcoal using a wet oxidation, stepped-combustion
procedure. Radiocarbon 42(2): 127-140.
Douka K, Hedges RM, Higham TG (2010) Improved AMS 14C dating of shell carbonates using high-precision X-ray diffraction and anovel density separation protocol (CarDS). Radiocarbon 52(2): 735-751.

Timing of the RNMH in Europe

Replacement process of Neanderthals by Modern Humans in
EuropeKatsuhiro Sano1* and Takayuki Omori1

1 The University Museum, The University of Tokyo, Japan. *Email: sano@um.u-tokyo.ac.jp

Since 2010, the RNMH project has been compiling an extensive archaeological database (Neander DB), which
covers the chronological period between c. 200 and 20 ka, spanning from Africa and across Europe, to Siberia,West, South and East Asia, and into Oceania (Nishiaki 2012; Sano et al. 2013). The European database contains
the richest records, including a total of 737 sites, 2,154 cultural horizons, and 3,453 radiometric dates (as of July 30,2014). In order to accurately reconstruct the process underlying the demise of Neanderthals and Modern Humandispersals, a rigorous examination of radiometric dates by advanced assessment of data reliability and Bayensian
analysis was undertaken.
Although previous studies reconstructed the gradual decline of Late Mousterian sites by 28 ka (van Andel et al.
2003) or suggested the late survival of Neanderthals in southern Iberia (Finlayson et al. 2006), our examination
indicates more rapid decrease of Neanderthal sites after Heinrich Event 5 (HE5). Unlike the recovery of the Late
Mousterian sites after MIS4, Neanderthals seem to have never increased their population size after HE5.
During the declination process of Neanderthal populations, the first Modern Human groups represented by the
Bachokirian/Bohunician industries expanded into Central Europe at c. 47 k cal BP, which coincides with abrupt
warming after HE5. As the number of the Bachokirian/Bohunician industries is relatively small, their population
size was probably not extremely large and it is uncertain whether or not the Bachokirian/Bohunician people further
migrated into Western and Southwestern Europe. Slightly after the first Modern Human dispersal, the Uluzzian
emerged in the Italian Peninsula by 45 k cal BP and the Chatelperronian at around 45 k cal BP in the FrancoCantabrian. Then, the Proto-Aurignacian appeared in the Mediterranean region at c. 43 k cal BP and expanded theterritory during short-period of time. Although the climate has dramatically changed between 45 k and 40 k cal BP
due to the Dansgaard-Oeschger events, the Proto-Aurignacian Modern Humans developed their population size.
In contrast, the population density of Neanderthals has extremely been reduced during this period and they became
extinct at latest during HE4.
References
Finlayson C, Giles Pacheco F, Rodríguez-Vidal J, Fa DA, Maria Gutierrez Lopez J, Santiago Perez A, Finlayson G, Allue E, Baena
Preysler J, Caceres I, Carrion JS, Fernandez Jalvo Y, Gleed-Owen CP, Jimenez Espejo FJ, Lopez P, Antonio Lopez Saez J, Antonio
Riquelme Cantal J, Sanchez Marco A, Giles Guzman F, Brown K, Fuentes N, Valarino CA, Villalpando A, Stringer CB, Martinez Ruiz
F, Sakamoto T (2006) Late survival of Neanderthals at the southernmost extreme of Europe. Nature 443: 850–853.
Nishiaki Y (2012) The archaeological database for the RNMH Project. In: Akazawa T, Nishiaki Y (eds) RNMH 2012 -The firstinternational conference on the replacement of Neanderthals by Modern Humans: Testing evolutionary models of learning. RHMH
Project Group, Tokyo, pp. 92–93.
Sano K, Kadowaki S, Naganuma M, Kondo Y, Shimogama K, Nagai K, Nakata H, Omori T, Yoneda M, Nishiaki Y (2013) Modern
Human dispersal into Eurasia: Preliminary results of the multi-disciplinary project on the replacement of Neanderthals by Modern
Humans (RNMH). In: Presented at the proceedings of European Society for the Study of Human Evolution 2, Leipzig, p. 195.
van Andel T, Davies W, Weninger B (2003) The human presence in Europe during the Last Glacial Period I: Human migrations andthe changing climate. In: van Andel T, Davies W (eds) Neanderthals and Modern Humans in the European landscape during the Last
Glaciation: Archaeological results of the Stage 3 Project. McDonald Institute for Archeological Research, Cambridge, pp. 31–56.

067
Timing of the RNMH in EuropeDiffusion or progressive integration? Re-considering thewidespread distributions of early UP/LSA bladelet technologies
from west Asia to south Europe and from east Africa to south Asia
Seiji Kadowaki

Nagoya University Museum, Nagoya University, Japan. Email: kadowaki@num.nagoya-u.ac.jp

Existing hypotheses for the widespread distributions of early UP bladelet technologies
The Late Pleistocene Out-of-Africa models of Homo sapiens, as variously delineated by genetic andpalaeoanthropological studies, have greatly influenced interpretations of behavioral and cultural records from
the Middle and Initial/Early Upper Palaeolithic periods in Eurasia. For example, several archaeologists haveproposed that lithic techno-typological similarity among some of the Initial Upper Palaeolithic or “transitional”
industries in the Levant (i.e., the Emiran), central Europe (i.e., the Bohunician), and the Altai (i.e., Kara Bom)can be regarded as “road-signs” of Homo sapiens groups expanding their geographic distributions from theLevant to other parts of Eurasia. Similarly, recently increasing Middle Palaeolithic (MP) records in the Arabian
Peninsula have greatly contributed to discussions on the early dispersal of Homo sapiens along the “southernroute”. Although these two cases of archaeological interpretations remain to be attested by human fossil records,
later widespread distributions of bladelet technologies during the Early Upper Palaeolithic (EUP) in westernand southern Eurasia are more widely accepted as cultural phenomena associated with Homo sapiens’ geographic
expansion. For example, several researchers suggested a technological and chronological correspondence between
assemblages under the category of the Ahmarian in the Levant and the Protoaurignacian in southern Europe.
These industries have been also compared with the Baradostian in the Zagros and EUP bladelet assemblages in
the Caucasus and discussed in relation to the model of Homo sapiens’ dispersal along the “northern route”. For
researchers who accept the hypothesis of earlier dispersal indicated by the Emiran-Bohunician cultural link, theAhmarian-Protoaurignacian connection means another dispersal event, and these multiple dispersal events are
considered to represent the “stepwise” colonization of Europe by Homo sapiens (Hublin 2013). On the other hand,the “southern route” hypothesis has also been archaeologically examined by comparing geometric microliths andsome decorative objects between late MSA/early LSA in east/south Africa and early UP in south Asia.

068

From a standpoint of the Out-of-Africa models, these widespread occurrences of bladelet technology havebeen interpreted to represent cultural diffusions from the Levant and Africa driven by population movement.
More precisely, the bladelet technologies are considered to have initially occurred in limited geographic areas,i.e., the Levant and Africa, and spread afterwards in association with the dispersal of Homo sapiens populations.
Such diffusion hypotheses have implications for learning behaviors involved in the spread of this technology
because the hypotheses distinguish between core areas, where technological/cultural innovations occurred (BarYosef 2007), and surrounding areas, where migrating groups from the core areas kept the same technology/
culture through social learning without further innovations although they probably faced new natural and social
environments. On the other hand, Le Brun-Ricalens et al. (2009) have proposed an alternative interpretation
for the widespread occurrences of bladelet assemblages in southern Europe and west Asia, suggesting theindependent mosaic appearance of bladelet technologies in various areas and their progressive integrations
through time. Instead of an innovation center, this view assumes indigenous technological developments in
multiple locations that were progressively connected with each other possibly through a technological transfer.
This paper considers these alternative interpretations for the wide geographic occurrences of EUP bladeletassemblages in western/southern Eurasia and east Africa by introducing two relevant archaeological records
investigated by Japanese expeditions from The University of Tokyo and Nagoya University.

Global Processes of RNMH

Wadi Kharar 16R and Mtongwe: Japanese investigations of early UP/LSA bladelet industries in
Syria and Kenya
An EUP site of Wadi Kharar 16R, located in the middle Euphrates, was discovered in archaeological survey
conducted by the Syria-Japan joint project between 2008 and 2011 (Nishiaki et al. 2012). We recorded more than
350 locations where artifacts or features (mostly Bronze Age tomb mounds) were found on the surface. The openair site of 16R is located on the left bank of Wadi Kharar. The site is comprised of two concentrations of lithics
on the sloping surface; a concentration in Area 1 measures ca. 10 m x 10 m and another in Area 2 is ca. 10 m x 5
m. Techno-typological characteristics of the chipped-stones excavated from this site are similar to those of Ksar
Akil Phase 4, which is typologically characterized by higher proportions of end-scrapers, particularly simple-end
scrapers. El-Wad points and retouched blades/bladelets are common and tend to be made on blanks with straightor curved, rather than twisted, profiles. Burins are predominantly dihedral types. Carinated pieces are present but
infrequent. Technologically, cores are dominated by single platform bladelet varieties, and blades/bladelets havestraight or curved profiles instead of twisted ones. Bladelets were produced from multiple strategies. Notably,
these techno-typological characteristics of Ksar Akil Phase 4 have recently been suggested to show strong
similarity to “the Early Ahmarian of the marginal zone” (Williams and Bergman 2010: 144) despite its former
categorization to the “Levantine Aurignacian”.
A Stone Age site of Mtongwe is located near the Kenyan coast, ca. 4 km south to Mombasa. It is an open-air
site-complex, where ca. 70 sites have been discovered in an area of ca. 800 m x 300 m on the marine terrace thatwas formed on the Jurassic shale covered by 30 m-thick Pleistocene deposits. The investigations of site started in1975 as part of Nagoya University East African Archaeological Prehistoric Research, followed by seven seasonsof fieldwork until 1989. The results of the excavations have been published in a series of reports (Omi 1977, 1980,
1982, 1984, 1986, 1988, 1991) although they have been rarely cited in the archaeological literature on African
MSA/LSA (except for Tryon and Faith 2013). The investigators defined three different lithic industries on thebasis of techno-typological features of the excavated lithic assemblages and their stratigraphic sequences. The
first industry, characterized by handaxes, large flakes, and side scrapers, has been defined by assemblages fromthe lowest gravel layers immediately above the bedrock. The above, second industry has been defined by lithic
assemblages including small bifaces, Levallois cores/flakes, and discoidal cores from upper sand-silt deposits. The
third industry, relevant to this paper, includes lithic assemblages from red sand deposits that cover the top of themarine terrace where the site is located, and it is notably characterized by the occurrences of Levallois debitage,
backed bladelets, and geometric microliths made on bladelets detached from single-platform prismatic cores. The
proposed association of Levallois and bladelet products in the same assemblage is shown by piece-plotted ltihic
distributions in four cultural layers. On the basis of such spatial/stratigraphic analyses, the investigators identified
three stratified sub-groups that show gradual increase in the proportions of microlithic components as opposed to
the Levallois debitage.
Drawing on the above new and little known data, the paper discusses issues on the timing and the process for the
emergence of bladelet technologies in the Levant and East Africa. The discussions also include their implications
for the existing hypotheses for the widespread occurrences of bladelet technologies in southern Europe and south
Asia.
References
Bar-Yosef O (2007) The archaeological framework of the Upper Paleolithic Revolution. Diogenes 54(3): 3-18.
Hublin J-J (2013) The makers of the early Upper Paleolithic in western Eurasia. In: Smith F, Ahern J (eds) The origins of Modern
Humans: Biology reconsidered. John Wiley & Sons, Inc., New Jersey, pp. 223-252.
Le Brun-Ricalens F, Bordes J-G, Eizenberg L (2009) A crossed-glance between southern European and Middle-Near Eastern earlyUpper Palaeolithic lithic technocomplexes: Existing models, new perspectives. Camps M, Szmidt C (eds) The Mediterranean from
50000 to 25000 BP: Turning points and new directions. Oxbow Books, Oxford, pp. 11-33.
Nishiaki Y, Sultan A, Kadowaki S, Kume S, Shimogama S (2012) Archaeological survey around Tell Ghanem al-‘Ali (V). Al-Rāfidān
33: 1-6.

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Omi G (1977) Third preliminary report of African studies (Archaeology 1). Association of African Studies, Nagoya University.
Omi G (1980) Fifth preliminary report of African studies (Archaeology 2). Association of African Studies, Nagoya University.
Omi G (1982) Mtongwe 1980: Seventh preliminary report of African studies (Archaeology 3). Association of African Studies,
Nagoya University.
Omi G (1984) Mtongwe 1982: An interim report of the East and Northeast African Prehistory Research Project 1982. Shinshu
University.
Omi G (1986) Mtongwe 1984: An interim report of the East and Northeast African Prehistory Research Project 1984. Shinshu
University.
Omi G (1988) Mtongwe and Mgonga: An interim report of the East and Northeast African Prehistory Research Project 1986.
Shinshu University.
Omi G (1991) An interim report of the East and Northeast African Prehistory Research Project 1988 and 1989. Shinshu University.
Tryon CA, Faith JT (2013) Variability in the Middle Stone Age of eastern Africa. Current Anthropology 54, Supplement 8:
S234-S254.
Williams J, Bergman C (2010) Upper Paleolithic levels XIII-VI (A and B) from the 1937-1938 and 1947-1948 Boston College
excavations and the Levantine Aurignacian at Ksar Akil, Lebanon. Paléorient 36(2): 117-161.

Fig. 1 Chipped stones (retouched tools) from an EUP site of Wadi Kharar 16R
in the middle Euphrates, northern Syria.

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Fig. 2 Geometric microliths from a late MSA/early LSA site at Mtongwe, east
Kenya.

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Neanderthals and Modern Humans in the Indus Valley(Sindh, Pakistan): The Middle and Late Palaeolithic of aforgotten region of the Indian Subcontinent
Paolo Biagi1* and Elisabetta Starnini2

1. Dipartmento di Studi sull’Asia e sull’Africa Mediterranea, Università Ca’Foscari Venezia, Italy; 2. Dipartmento di Studi Storici, Scuola
di Scienze Umanistiche, Università Degli Studi di Torino, Italy. *Email: pavelius@unive.it

This paper describes and discusses the Middle and Late (Upper) Palaeolithic sites of Sindh (Pakistan), a regionof the Indian Subcontinent whose early prehistory, except for the Bronze Age Indus Civilization, has never beentaken into great consideration by most archaeologists.
Sindh, and the Indus Valley in general, are territories of basic importance for understanding the spread of bothNeanderthals and first Modern Humans towards south-east Asia. Although some authors suggested that the firstModern Humans moved along the northern coast of the Arabian Sea, crossing both present Las Bela (Balochistan)and Lower Sindh on their way to India, no evidence of their passage has so far been recorded.
Most of the Middle Palaeolithic industries and tools so far recorded from the above regions come from thealmost unpublished geological surveys made in the 1970’s by Professor A.R. Khan in Lower Sindh, and the brief
visits by B. Allchin in Upper Sindh. More discoveries were made during the last 30 years mainly at Ongar, near
Hyderabad (Lower Sindh), and in the Rohri Hills, near Sukkur (Upper Sindh), by the Italian Archaeological
Mission.
The presence of characteristic Levallois Mousterian assemblages at Ongar (Lower Sindh), and other sites along
the southwestern bank of the Indus River, opens new perspectives for the study of the dispersal of Middle
Palaeolithic Neanderthals, whose south-easternmost spread has systematically been neglected by most authors.
Although the presence of typical Levallois Mousterian assemblages has been recorded from Iran, Afghanistan,
Uzbekistan, and former Soviet Central Asia in general, evidence of the above complexes in the Indian
Subcontinent is very poor. In effect the occurrence of typical Levallois discoid cores, flakes, blades, points,
Mousterian scrapers and one Mousterian point at Ongar seem to mark the south-easternmost distribution of thiscultural aspect, which does not characterize the Middle Palaeolithic of the Indian Subcontinent, given that thislatter is mainly characterized by unretouched flake assemblages and scrapers. Nevertheless, Levallois points andflakes have already been described as one minor component of the so-called “Late Soan” complexes of Punjab
farther north along the same bank of the Indus.
Unfortunately, all the Levallois Mousterian assemblages so far recorded from Lower Sindh can be attributed to
the Middle Palaeolithic only on the basis of their typological and technological characteristics.. In fact they havebeen collected as surface find spots without associated organic material suitable for absolute dating.
Even more complex is the definition of the earliest Late (Upper) Palaeolithic assemblages in the study region.
In contrast with what previously suggested, Late (Upper) Palaeolithic sites are quite common in some areas ofLower Sindh, among which are the Mulri Hills (Karachi) and Jhimpir (Thatta). The assemblages from these sites
are characterised by subconical cores, bladelets, lunates of different shape and size, and, in a few cases, a highpercentage of Burins. Given their typological characteristics they seem to belong to quite an advanced stage inthe development of the Late (Upper) Palaeolithic period.

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The situation in Upper Sindh is quite different. The flat limestone terraces of the Rohri Hill yielded evidenceof an impressive number of Late (Upper) Palaeolithic flint workshops, characterised by subconical bladelet andbladelet-like flakelet cores, and millions of debitage products. A similar situation has been recorded also from
Ongar, even though most of the workshops from these hills have been devastated by recent industrial limestonemining.
To conclude: Sindh is an extremely important country for the study of the Palaeolithic of the Indian
Subcontinent and its related territories. It is unfortunate that, due to many factors, our knowledge of such animportant territory is far too poor.

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Global Processes of RNMHNew insight into the end of the Middle Palaeolithic in Central
Asia
Yoshihiro Nishiaki1*, Otabek Aripdjanov2, Rustam Suleymanov3, Hiroto Nakata4,
Saiji Arai5, Takehiro Miki5, Takayuki Omori1 and Jannat Ismailova2
1. The University Museum, The University of Tokyo, Japan; 2. The State Museum of the History of Uzbekistan, The Republic of
Uzbekistan; 3. National University of the Republic of Uzbekistan, Republic of Uzbekistan; 4. Faculty of Letters, Aoyama Gakuin
University, Japan; 5. Department of Archaeology, The University of Tokyo, Japan.
*Email: nishiaki@um.u-tokyo.ac.jp

Central Asia has become a focal region for investigating varied processes of the replacement of Neanderthals byModern Humans. These processes might have been more complicated than in West Eurasia, involving at least
one other hominin group (Denisovans). Recent multidisciplinary research in the Altai Mountains of eastern
Central Asia has revealed the occurrence of at least three different Middle Paleolithic lithic industries (Denisova,
Kara-Bom, and Sibiryachikha), likely maintained by different hominin populations (Derevianko et al. 2013).
Similarly, in western Central Asia, apparently different industries (Obi Rahmat and Teshik Tash) are known
in the Middle Palaeolithic (Derevianko 2010). Fossil records also indicate the possible presence of at least two
different hominin groups: Neanderthals (Teshik Tash) and populations displaying mixed traces of anatomically
Modern Humans and Neanderthals (Obi Rahmat; Viola et al. 2004; Glantz et al. 2008).
To interpret these findings in relation to the replacement of indigenous populations by the presumed incoming
Modern Humans, it is essential to establish a rigorously controlled chronological framework to which particular
sets of lithic industries and fossil records are assigned. However, reliable radiometric dates are available only at
limited sites (e.g., Denisova Cave and Obi Rahmat). To develop this framework, a larger array of dates fromsecurely stratified contexts is clearly needed.
Our prehistoric research at Anghilak Cave, southern Uzbekistan, aimed to provide new data on its Middle
Palaeolithic occupations, thereby contributing to construction of an adequate chronological framework. Locatedapproximately 50 km southwest of Samarkand, the site was previously excavated by an Uzbek-American team
between 2002 and 2004. A non-diagnostic human metatarsal was recovered in association with Mousterian
lithic assemblages from contexts dated to ca. 42–32 ka (Beeton et al. 2014), a rather late date for the Middle
Palaeolithic. Our fieldwork at this cave in 2013 and 2014 consisted of stratigraphic sampling of the archaeologicaland environmental materials as well as radiocarbon dating the Middle Palaeolithic deposits. Preliminary
results addressed in this paper indicate that the Middle Palaeolithic occupations of this cave ended earlier thanpreviously reported. Furthermore, the recovered lithic assemblages differ from contemporaneous assemblages
from Obi Rahmat, the site with the best Middle Palaeolithic stratigraphic sequence in Uzbekistan, suggesting theco-occurrence of different Mousterian industrial facies in western Central Asia.
References
Beeton TA, Glantz MM, Trainer AK, Temirbekov SS, Reich RM (2014) The fundamental hominin niche in late Pleistocene Central
Asia: A preliminary refugium model. Journal of Biogeography 41: 95–110.
Derevianko AP (2010) Three scenarios of the Middle to Upper Paleolithic transition. Scenario 1: The Middle to Upper Paleolithic
transition in Central Asia and the Near East. Archaeology, Ethnology and Anthropology of Eurasia 38(4): 2–38.
Derevianko AP, Markin SV, Shunkov MV (2013) The Sibiryachikha facies of the Middle Paleolithic of the Altai. Archaeology,
Ethnology and Anthropology of Eurasia 41(1): 89–103.
Glantz M, Viola B, Wrinn P, Chikisheva T, Derevianko AP, Krivoshapkin A, Islamov U, Suleimanov R, Ritzman T (2008) New
hominin remains from Uzbekistan. Journal of Human Evolution 55: 223–237.
Viola B, Seidler H, Nedden D (2004) Computer tomographic investigations of the OR-1 petrosals. In: Derevianko AP (ed) Obi-

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Rakhmat Grotto. Institute for Archaeology and Ethnography, Novosibirsk, pp.100–106.

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The exploitation and adaptation in the Asian Arctic andPaleolithic in the Baikal Siberia
Ekaterina A. Lipnina1, German I. Medvedev1, Fedora Khenzkhenova2,
Kunio Yoshida3, Dai Kunikida3, Takao Sato4, Kenji Suzuki5and Hirofumi Kato5*
1. Irkutsk State University, Russia; 2. Siberian Branch of Russian Academy of Science, Russia; 3. The University of Tokyo, Japan; 4. Keio
University, Japan; 5. Hokkaido University, Japan. *Email: h-kato@let.hokudai.ac.jp

The exploitation and adaptation in the Arctic zone by Modern Human is one of the significance issues of humanhistory. Recent studies in the Arctic area demonstrated that human activities in the polar regions of 70°N is
traced back to 40,000 years old (Pitluko et al. 2012; Питулько и Павлова 2010). However, it is still not clear their
origin and techno-cultural basis enabled human group to adapt to the Arctic environment. In order to explore theorigin of the pioneers of the first, it is necessary to clarify Early Upper Paleolithic populations in southern Siberia
and their technological basement.
In this report, we would like to present the key-points to consider the dispersal and adaptation behavior byModern Human in the MIS3, based on the results of our joint researches focused on the transition period
between Middle to Upper Paleolithic in the Baikal Siberia. The Middle Paleolithic in Eastern Siberia had
been characterized original characteristics without typical levallois reduction technique. And Upper Paleolithic
industries, has evolved technically through the Middle to Upper Paleolithic transition subsequent spread from
Baikal Siberia to northern Eurasia, including the Arctic zone. Technological characteristics of lithic industries
in the Middle to Upper Paleolithic transition period characterized by the emergence of systematic microblade
reduction. With this new innovation, Modern human has become successful development and adaptation to the
new living environment.
We have continued to explore lithic industries of data MIS3 in Baikal Siberia as the Japanese-Russian joint
research from 2005. Through this joint research, We were able to provide cultural sequence from Middle to Upper
Paleolithic accompanied by faunal remains beside of the Angara River. In Borl’shoi-Nalin 1 site, one of the survey
sites in this river basin, was discovered remains of long-term human activity in the cut bank wall and as beachdeposits in the modern floodplain.
Bones and Charcoals on the cultural layer from excavation area have been direct-dated between 28,000-31,000
yrBP and 40,000-41,000 yrBP. It has the potential to be confirmed cultural layer to continue more lower partof deposit. From the results of excavation from Bol’shoi-Nalin 1 site, suggest that the lithic industries stableassociated with microlithic, had widely spread around of this area in the second stage of MIS3.
The lithic industry of the Early Upper Paleolithic in Siberia, are known different technological traditions. These
technological diversity of lithic industries represents the reflection of the diversity of the human groups at thattime. Also, the characteristics seen on their cultural-technological traditions and of their artworks prospect usthat there had the immigrants from Western Siberia and Russia plain, which means the human evolution of thisarea was unilineal one. Therefore, we could assume that there were various and intricate human evolution stream
in Eastern Siberia Area.
We could say that Middle to Upper Paleolithic transition in Eastern Siberia provide us the very valuable
materials from which we can see broadness of the Anatomical human being and their culture in this area, andalso the diversity of these formation process. As a turning point in the development of the Arctic area, human075

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groups could expand their activity spaces dramatically even further more to the new continent.
References
Pitulko VV, Pavlova EY, Nikolskiy PA, Ivanva VV (2012) The oldest art of the Eurasian Arctic: Personal ornaments and symbolic
objects from Yana RHS, Arctic Siberia. Antiquity 86 (333): 642-659.
Питулько ВВ, Павлова ЕЮ (2010) Геоархеология и радиоуглеродная хронология камменного века Северо-Восточной Азии.
Наука, Санкт-Петербург.
Sato T, Khenzkhenova F, Simakova A, Danukolova G, Morosova E, Yoshida K, Kunikita D, Kato H, Suzuki K, Lipunina E,
Medvedev G, Martynovich N (2014) Paleoenvironment of the Fore-Baikal region in the Karginian interstadial: Results of theinterdisciplinary studies of the Bol'shoj Naryn site. Quaternary International 333: 146-155.
Khenzkhenova F, Sato T, Lipunina E, Medvedev G, Kato H, Kogai S, Maximenko K, Novosel'zeva V (2011) Upper Paleolithic
mammal fauna of the Baikal region, east Siberia (new data). Quaternary International 231: 50-54.
Kato H (2013) The Middle to Upper Paleolithic transition in Siberia: Three regional sketches for replacement. In: Akazawa T,
Nishiaki Y, Aoki K (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 1: Cultural perspectives. Springer, New
York, pp. 93-103.

076
Global Processes of RNMHInferring possible contacts between Denisovan and someancestors of modern East Eurasians
Naruya Saitou1,2*, Timothy A. Jinam1,2 and Hideaki Kanzawa-Kiriyama2,3

1. Division of Population Genetics, National Institute of Genetics, Japan; 2. Department of Genetics, Graduate University for Advanced
Studies, Japan; 3. Department of Anthropology, National Museum of Nature and Science, Japan. *Email: saitounr@nig.ac.jp

Study of human populations on Japanese Archipelago
Japanese Archipelago stretches over 4000 km from north to south, and is homeland of three human populations;
Ainus, Main-islanders, and Ryukyuans. Origins of these people have been studied for long time. Standard
theory based on craniofacial data is “dual structure model”. According to this model, first migrants to JapaneseArchipelago came from somewhere in Southeast Asia in Upper Paleolithic age, who were ancestors of Jomon
people. Second wave of migration took place later in Yayoi period (3000 BP to 1700 BP), and people came in thistime from Northeast Asia. Indigenous Jomon people and new migrants in and after Yayoi period gradually mixed
with each other. This model provides reasonable explanation for morphological similarity between Ainu people of
Hokkaido, northernmost main island of Japanese Archipelago, and Ryukyuan people in Southwest Archipelago,
despite of large geographical distance. Japanese Archipelago Human Population Genetics Consortium (2012)
determined genome-wide 0.9 million SNPs for Ainus and Ryukyuans, and compared these with existingdata sets. This was first report of these genome-wide SNP data, and the dual structure model was essentiallyconfirmed. We also examined nuclear genome sequences of 3,000 year-old Jomon people whose remains were
found from Sanganji Shell Mound (Fukushima Prefectural Museum 1988), using DNA extracted by KanzawaKiriyama et al. (2013) for mitochondrial haplotype analyses (Kanzawa-Kiriyama et al. unpublished). We foundthat Jomon people were very different from modern East Eurasians (Li et al. 2008; The 1000 Genome Project
Consortium 2012), and they are probably basal to all presentday East Eurasians. Furthermore, some gene flowfrom Denisovan genome (Reich et al. 2010; Meyer et al. 2012) to Jomon ancestors was suspected throughstatistical analyses as well as that from Neandelthal genome (Green et al. 2010).
Study of Negrito populations on South East Asia
Negritos distributed in Philippines Islands, Malay Peninsula, and Andaman Islands are considered to be remnantsof early human migrations to Sundaland. Jinam et al. (2012) determined mitochondrial genome sequences for
86 human individuals including Malaysian Negritos, and conducted phylogenetic analysis. They also reanalyzed
genomewide SNP data of people in South East Asia reported by HUGO Pan-Asian SNP Consortium (2009).
We found that Malaysian and Philippine Negritos were quite different from other South East Asian populations.
We proceeded to analyze close to one million genomewide SNP data of Malaysian, Philippine, and Andaman
Negritos, and found some similarity among them. When we compared these data with Denisovan genome
data, Philippine Aeta Negritos of Luzon Island showed higher gene flow from Denisovans than Mamanwaof Mindanao Island. This finding further strengthens hypothesis that Denisovan people admixed with earlymigrants from Africa probably somewhere in Sundaland.
References
HUGO Pan-Asian SNP Consortium (2009) Mapping human genetic diversity in Asia. Science 326: 1541–1545.
Japanese Archipelago Human Population Genetics Consortium (2012) The history of human populations in the Japanese Archipelago inferred from genome-wide SNP data with a special reference to the Ainu and the Ryukyuan populations. Journal of077

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Human Genetics 57: 787-795.
Kanzawa-Kiriyama H, Saso A, Suwa G, Saitou N (2013) Ancient mitochondrial DNA sequences of Jomon teeth samples from
Sanganji, Tohoku district, Japan. Anthropological Science 121: 89-103.
Reich D, Green RE, Kircher M, Krause J, Patterson N, Durand EY, Viola B, Briggs AW, Stenzel U, Johnson PL et al (2010)
Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature 468: 1053-1060.
Fukushima Prefectural Museum (ed) (1988) Sanganji Shell Mound. Fukushima Prefectural Museum, Fukushima. (In Japanese)
Jinam TA, Hong LC, Phipps MA, Stoneking M, Ameen M, Edo J, Pan-Asian SNP Consortium, Saitou N (2012) Evolutionary history of continental South East Asians: “Early train” hypothesis based on genetic analysis of mitochondrial and autosomal
DNA data. Molecular Biology and Evolution 29: 3513-3527.
Meyer M et al (2012) High-coverage genome sequence from an archaic Denisovan individual. Science 338: 222-226.
Green RE et al (2010) A draft sequence of the Neandertal genome. Science 328: 710-722.
The 1000 Genome Project Consortium (2012) An integrated map of genetic variation from 1,092 human genomes. Nature 491:
56-65.
Li JZ, Absher DM, Tang H, Southwick AM, Casto AM, Ramachandran S, Cann HM, Barsh GS, Feldman M, Cavalli-Sforza
LL et al (2008) Worldwide human relationships inferred from genome-wide patterns of variation. Science 319: 1100-1104.

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DAY 5: December 4, 2014

Learning Strategies and Cultural Evolution

Lives to learn? The biology and culture of Neanderthal andearly Modern Human learning behaviours
Olaf Jöris

MONREPOS Archaeological Research Centre and Museum for Human Behavioural Evolution, Germany. Email: joeris@rgzm.de

The human brain is organized entirely different from that of any other primate, implying significant disparity incognitive abilities. The human brain also seems to be more sensitive to evolutionary changes, such as mutations,which could have affected large parts of our brains during our evolutionary history. In addition, human lifehistories differ much from those of any other primate. Here it will be discussed, first, whether or not and towhich degree and extent such modern-day observations can be transferred back into the past and – second –what these observations imply for early hominin learning behaviours. The implications for the reconstruction of
past lifeways will be considered and compared with the Middle Stone Age (MSA) and Middle Palaeolithic (MP)archaeological records. Special focus will be on data on land-use strategies, regional cultural variation (RCV) andtool production and standardization. Similarities and differences between these records and their implications forunderstanding possible differences in the modes of learning of Neanderthals and early Modern Humans will be
evaluated.

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TTT: Tradition, (archaeological) taxonomy and (cultural)transmission
Anna Belfer-Cohen1* and Erella Hovers1

1. Institute of Archaeology, The Hebrew University of Jerusalem, Israel. *Email: belferac@mscc.huji.ac.il

Our perception of the prehistoric past is formulated by material culture taxonomies. These taxonomies originate
through processes of archaeological exploration. Because the taxonomies are not natural constructs, they are basedon particular circumstances, specific phenomena, or loosely defined, broad criteria mixing chronology, geography,
techno-typology or any of their combinations. ‘Initial Upper Paleolithic’, ‘Mousterian of the Acheulian Tradition’,‘The Ahmarian’ are examples of such fluid taxonomies. Therefore, there is an inherent flaw in the endeavors tointerpret data within such a framework, culminating in self-fulfilling prophecies.
How can archaeologists untie this Gordian knot? A first step has already been taken – prehistorians do not speakof cultures but of different scales of the archaeological phenomena (for example, techno-complexes), i.e., they
adhere more closely to the factual record, attempting to minimize the broad (and often vague) connotations
to definitions of extant ‘cultures’. Another caveat to bear in mind is that there is a discrepancy between theindividual human time frame, within which behavioral decisions are made, and the evolutionary time scale ofarchaeological taxonomies, which time-averages many such episodes of decision-making.
With these epistemological difficulties in mind, we explore in this paper the relationship between behavioral
processes (“knowledge transmission”) and their long-term outcomes, among which is the formation of prehistoric“traditions”, trying to identify their respective imprints within the prehistoric record.

081
Learning Strategies and Cultural Evolution – Archaeological ApproachCultural transmission in the Pleistocene: Studying
Neanderthal-Modern Human contact in Central Europe
during OIS3
Gilbert B. Tostevin

University of Minnesota, USA. Email: toste003@umn.edu

If we want to differentiate among the many hypotheses for why anatomically Modern Humans from Africa
outcompeted and replaced evolutionary relatives in Eurasia such as the Neanderthals, it is necessary to
identify the geographical and adaptive contexts in which the contact between these different homininsproduced different results. If the Modern Humans immediately replaced Neanderthals in one context but in
another context remained in competition with Neanderthals or influenced Neanderthal cultural behavior fora period of time, the comparison of the behavioral details between these two contexts should highlight whatgave Modern Humans the greater advantage. The comparison of the archaeological records of the Middle
to Upper Paleolithic (MP-UP) transition between Western Europe and Central Europe could do exactly
this. For instance, the transition began at least 5 ka earlier in the Middle Danube of Central Europe than inWestern Europe (Müller et al. 2011) and produced a greater diversity of industrial types, with two “transitional”
industries (the Bohunician and the Szeletian) rather than one, as with the Châtelperronian in France and theUluzzian in Italy. Yet despite their early appearance, the two “transitional” industries of Central Europe did
not spread as widely or last as long as the subsequent Aurignacian. Understanding why the latter industry
went “viral” across western Eurasia while the former industries did not will help us understand why Modern
Humans survived the Pleistocene and our evolutionary relatives did not.
The present paper argues that studying the cultural transmission processes involved in the MP-UP transitionin Central vs. Western Europe is critical to understanding why Modern Humans were able to replace
Neanderthals. Yet such a study will only make progress if archaeologists are able to analyze lithic assemblages(and other material culture records as well) using methods designed from first principles to fit the question,i.e., to be sensitive to cultural transmission processes (sensu Boyd and Richerson 1985) according to applicablemiddle-range theory predictions for how specific material culture production behaviors are learnable atdifferent degrees of social intimacy (Tostevin 2007, 2012). Instead of studying traditional retouch typologicalcategories or chaîne opératoire-defined industries (what Shea 2014 calls named stone tool industries or“NASTIES”), the approach advocated here produces quantitative measures of similarity vs. dissimilarity
between assemblages using variables that have empirically-verifiable relationships to the social transmissionof technical knowledge, i.e., the learning process itself.

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The present paper makes the case for how this Behavioral Approach to Cultural Transmission (BACT)
can improve our study of the Middle to Upper Paleolithic transition. It does this through a comparisonof the respective abilities of the BACT method vs. the traditional approach to Paleolithic systematics totest scientif ically two hypotheses concerning the role of cultural transmission processes during the MPUP transition in Central Europe. There are currently two alternative hypotheses for the similarity in thelithic artifacts from industries attributed to the Bohunician in Central Europe with those of the Initial
Upper Paleolithic (IUP) assemblages of the Levant. In the first hypothesis, Tostevin (2003a, 2003b, 2012)
applied the attribute analysis tools of the BACT method to 19 assemblages in the Levant, Central Europe,and Eastern Europe to test the continuity vs. discontinuity in blank production behaviors and tool kit
morphologies between 60 and 30 ka 14C BP. In each region, the pattern of change is marked by an identicalsuite of flintknapping behaviors which had no precedent in any of the three regions. The chronogeographic

Learning Strategies and Cultural Evolution – Archaeological Approachpattern of this data, labeled the “Bohunician Behavioral Package (BBP)”, is consistent with the predictionsof a cultural transmission event from the Levant, through Southeastern Europe, and ending in Central
and Eastern Europe. The second hypothesis is Zilhão’s (2006) suggestion that the similarities between theEuropean Bohunician and Levantine IUP assemblages are coincidental and that the Bohunician represents
a cultural continuity across the MP-UP transition. Using TL dates, artifact illustrations, and the chaîne
opératoire descriptions of the assemblages from Piekary IIa, Poland (Sitlivy et al. 1999; Valladas et al. 2003),
Zilhão concludes that the southern Polish assemblages represent “the in-situ development of volumetric
Upper Paleolithic methods of blade debitage out of Levallois flake-based technologies” (2006: 187,189)
and thus are geographically more parsimonious candidates for antecedents of the Bohunician in the Middle
Danube Basin than the IUP of the Levant.
The present study contrasts how the BACT and traditional Paleolithic systematics evaluate these two
hypotheses. For the use of the BACT method for testing Zilhão’s hypothesis, the paper presents Tostevin’s
attribute analysis of the artifacts from Piekary IIa Layers 6, 7a, 7b, and 7c from the excavations conductedfrom 1998 through the early 2000s (trenches XX and XXII; Sitlivy et al. 2008). Using measures of similarity
vs. dissimilarity in blank production behaviors and tool kit morphologies between the Piekary IIa assemblages
and the BBP assemblages published to date, the results of the chronological pair-wise comparisons between
these assemblages are compared to the predictions from Zilhão’s hypothesis and those of the BBP hypothesis.
For the use of traditional Paleolithic systematics to test the two competing hypotheses, the paper evaluates
the arguments and data presented in Zilhão (2006) and Sitlivy et al. (2008). The successful design of futurestudies of cultural transmission in the Pleistocene are discussed in light of the advantages and disadvantages
revealed through the methodological comparison.
References
Boyd R, Richerson PJ (1985) Culture and the evolutionary process. University of Chicago Press, Chicago.
Müller U, Pross J, Tzedakis P et al (2011) The role of climate in the spread of Modern Humans into Europe. Quaternary Science
Reviews 30: 273-279.
Shea J (2014) Sink the Mousterian? Named stone tool industries (NASTIES) as obstacles to investigating hominin evolutionary
relationships in the Later Middle Paleolithic Levant. Quaternary International (http://dx.doi.org/10.1016/j.quaint.2014.01.024).
Sitlivy V, Zięba A (2006) Eastern and Central Europe before 30 kyr BP: Mousterian, Levallois and blade industries. In: Chabai V,
Richter J, Uthmeier T (eds) Kabazi II: The 70 000 years since the Last Interglacial. Shlyakh, Simferopol – Cologne, pp. 361-419.
Sitlivy V, Sobczyk K, Morawski W et al (1999). Piekary IIa Palaeolithic industries: Preliminary results of new multidisciplinary
investigations. Préhistoire Européenne 15: 45-64.
Sitlivy V, Zięba A, Sobczyk K eds (2008) Middle and Early Upper Palaeolithic of the Krakow region: Piekary IIa. Musées royauxd’Art et d’Histoire, Bruxelles.
Tostevin G (2003a) A quest for antecedents: A comparison of the Terminal Middle Paleolithic and Early Upper Paleolithic of theLevant. In: Goring-Morris N, Belfer-Cohen A (eds) More than meets the eye: Studies on Upper Palaeolithic diversity in the Near
East. Oxbow Press, Oxford, pp. 54-67.
Tostevin G (2003b) Attribute analysis of the lithic technologies of Stránská skála II-III in their regional and inter-regional context.
In: Svoboda J, Bar-Yosef O (eds) Stránská Skála: Origins of the Upper Paleolithic in the Brno Basin. Peabody Museum, Harvard
University, Cambridge, pp. 77-118.
Tostevin G (2007) Social intimacy, artefact visibility, and acculturation models of Neanderthal-Modern Human interaction. In:
Mellars P, Boyle K, Bar-Yosef O, Stringer C (eds) Rethinking the human revolution: New behavioural and biological perspectives on
the origins and dispersal of Modern Humans. MacDonald Institute for Archaeological Research, Cambridge, pp. 341-357.
Tostevin G (2012) Seeing lithics: A Middle-Range Theory for testing for cultural transmission in the Pleistocene. Peabody Museum,Harvard University, Cambridge, and Oxbow Books, Oxford/ Oakville.
Valladas H, Mercier N, Escutenaire C et al (2003) The Late Middle Paleolithic blade technologies and the transition to the Upper
Paleolithic in southern Poland: TL dating contribution. Eurasian Prehistory 1(1): 57-82.
Zilhão J (2006) Neandertals and Moderns mixed, and it matters. Evolutionary Anthropology 15: 183–195.

083
Learning Strategies and Cultural Evolution – Archaeological ApproachTransculturation versus acculturation: A clarification

Foni Le Brun-Ricalens

National Archaeological Research Center of Luxemburg, Great-Duchy of Luxemburg. Email: Foni.LeBrun@cnra.etat.lu

Several models have been proposed to explain processes underlying the transition between different technocultural assemblages in prehistoric archaeology. These ‘transitions’ either represent phenomena of ‘gradualism’
connected to in situ evolution or ‘diffusionism’ by various ‘acculturation’ processes prone to external influences(direct loans) and necessarily implicating long-distance migrations of populations. Following a review of theoriginal formulation of these two processes, an alternative paradigm is proposed – ‘transculturation’. Borrowed
from ethnologists and introduced by F. Ortiz in 1940, this process is characterised by the integration(through indigenous re-interpretation) of external influences via indirect loans derived from intimate
interpersonal contacts. In the sense of the term employed here, transculturation can take several different
forms (imitation, assimilation, hybridisation, re-interpretation) that are better suited to accounting for thediverse transformations evident in the archaeological record. Contrarily to acculturation which imposes new(foreign) manners of doing things, transculturation reinvests the people hidden behind each techno-culture
as the primary agents of their own transformation in that they may or may not be open to the diffusion ofcertain external ideas and have the possibility of re-interpreting them rather then suffering them.

084
Learning Strategies and Cultural Evolution – Archaeological ApproachToward an understanding of learning strategy in the Upper
Paleolithic: A case study in Hokkaido, Northern Japan
Jun Takakura

Archaeological Research Center, Hokkaido University, Japan. Email: jun-ta@let.hokudai.ac.jp

Introduction
The learning behaviors of Paleolithic hominins are currently one of the key questions in archaeological andpaleoanthropological research, in relation to development of the chaîne opératoire approach in the lithic
technological studies. Since the 1990’s, much attention has been focused on the skill learning behaviors of lithic
production at the Upper Paleolithic sites in Europe, in order to reveal the technological and social relationshipbetween experts and novices in lithic knappers (e.g. Pigeot et al. 1990). Recently I have demonstrated that thevarious learning behaviors for acquiring skill relevant to lithic production, based on the analyses of numerous
obsidian refitted artifacts, were conducted at the Upper Paleolithic sites in Northern Japan (Takakura 2013).
Importantly these researches have enabled us to know how we can elucidate skill learning behaviors fromthe archaeological record empirically, including meaningful applications of “expert”, “novice”, “instruction”,“exercise”, “pedagogic demonstration” and “trial and error” as a means of examining variability in the technicalskill levels among the lithic refitted artifacts. Although the detailed behavioral reconstructions relevant to theskill acquisition in the respective Paleolithic sites have been addressed, prehistoric archaeologists still understandlittle about how the learning behaviors in the past were actually related to the behavioral system of the prehistoric
hunter-gatherers such as mobility, territoriality, subsistence, settlement, and regional interaction. To emphasize
the behavioral and social contexts of lithic skill learning processes may lead in productive directions becausesuch an approach can provide us with important insights into situational and institutional variability of learning
behaviors in a given society.
The purpose of this paper is to show a case study on the skill learning strategies of the lithic knappers throughcomparative analysis of the late Upper Paleolithic assemblages in Hokkaido, Northern Japan. In this paper, I
consider this issue to integrate data about the learning behaviors for blade production and the updated knowledge
about inter-assemblage variability reflecting the overall behavioral system.
Learning behaviors and blade production in the late Upper Paleolithic
I focus on some of the microblade industries with the Hirosato type microblade cores from the Kamishirataki
2 (concentration Sb-9), Nittou, Takase, Ankaritou 7 sites, because these industries represent very specifictechnological features. Technologically identical blade and microblade reduction sequences are seen in these sites.
Large chunks of debris were commonly used as raw material of stone artifacts. Discovered formal and long blades(from almost 10 cm up to 30 cm in length) were subsequently detached with careful preparation and several
rejuvenation stages. We can often see rubbing on the pecked platform, presumably with the aid of abrasives. Suchblades were generally used as blanks of end-scrapers, side-scrapers, gravers, and the Hirosato type microbalde
cores. These industries belong to the late Upper Paleolithic according to the techno-typological perspective(Nakazawa et al. 2005).
The intensive manufacture of formal and long blades from blade cores was exclusively seen in the Kamishirataki

085

Learning Strategies and Cultural Evolution – Archaeological Approach2 site. Based on the analysis of technological characteristics, I suggested that four refitted sets relevant to the
manufacture of large blades from concentration Sb-9 at the Kamishirataki 2 site were produced by the highly
skilled knappers. It is of interest to note that many of the blades detached from the blade cores in these sets wereabandoned at concentration Sb-9 unlike the other numerous refitted sets in Shirataki. One conclusion that canbe drawn from this is that the expert knappers were not aiming at producing good blades for immediate use, butrather instructing beginners. These materials can be interpreted as “academic cores” (Takakura 2013).
On the other hand, at the Nittou, Takase, and Ankaritou 7 sites, the manufacture of microblades and several
rejuvenations of stone tools such as end-scrapers and gravers were only carried out. Stone tools of these sites
are dominated by numerous end-scrapers, side-scrapers, gravers, and graver spalls. In addition, hearths anddense of charcoals around the lithic concentrations were exclusively recovered from the Nittou and Ankaritou 7
sites. Consequently it is difficult to recognize traces of learning behaviors for blade production from these sites,
although various data in relation to daily and domestic activities can be apparently acquired.
Conclusion
The Kamishirataki 2 site is located near a huge outcrop of obsidian, whereas the Nittou, Takase, and Ankaritou
7 sites are located far from the raw material sources (from almost 30 km up to 180 km in distance). Thesesituations might lead to a difference in place and opportunity of the blade production and the learning behaviors
in the microblade industries with the Hirisato type microblade cores. The availability of raw materials probably
permitted such pedagogic demonstration by expert knappers.
The available data show that the concentration Sb-9 at the Kamishirataki 2 site was used specifically for the
manufacture of blades and bifacial points, and does not contain traces of daily and domestic activities such as
maintenance of stone tools. It is apparent that the formation of this concentration was related to the specificactivities by knappers responsible for the manufacture of stone tools. This result suggests that the learning
behaviors for blade production at the Kamishirataki 2 site were conducted by specific members away from thecontext of daily activities.
References
Nakazawa Y, Izuho M, Takakura J, Yamada S (2005) Toward an understanding of technological variability in microblade assemblagesin Hokkaido, Japan. Asian Perspectives 44: 276-292.
Pigeot N (1990) Technical and social actors: Flintknapping specialists at Magdalenian Etiolles. Archaeological Review from Cambridge 9: 126-141.
Takakura J (2013) Using lithic refitting to investigate the skill learning process: Lessons from Upper Paleolithic assemblages at the
Shirataki sites in Hokkaido, Northern Japan. In: Akazawa T, Nishiaki Y, Aoki K (eds) Dynamics of learning in Neanderthals andModern Humans, vol. 1: Cultural perspectives. Springer, New York, pp. 151-171.

086
Learning Strategies and Cultural Evolution – Archaeological ApproachFig. 1 Map showing the sites yielding the industries with the Hirosato type microblade cores.
Table 1 List of the number of stone artifacts in the industries with the Hirosato typemicroblade cores.

End-scraper
Side-scraper
GraverSpall
Drill
Bifacial point
Flake adze
Bifacial axe
Blade
Microblade
Blade core
Microblade core
Flake
Hammerstone
Grindstone

Sb-9, Kamishirataki 2
15
12
3
4
0
0
0
0
279
69
0
12
1340
0
0

Nittou
3
3
2
4
0
0
0
0
6
257
0
3
1067
0
0

Takase
15
11
28
65
6
5
4
0
65
245
1
8
25
0
0

Ankaritou 7
31
7
57
247
9
3
6
1
82
193
0
9
1426
3
1

087
Learning Strategies and Cultural Evolution – Archaeological Approach

Innovation, processes of social learning and modes ofcultural transmission amongst the Chabu adolescent foragerfarmers of Ethiopia
Bonnie Hewlett

Washington State University, Vancouver, Canada. Email: hewlettb@vancouver.wsu.edu

Innovation, as an element of behavioral plasticity, has been hypothesized to enhance the fitness andsurvivability of individuals, while overall increasing the diversity of cultural traits overtime. Based uponan earlier study of innovations and their transmission and acquisition amongst Aka forager adolescents ofcentral Africa, this study examines the same topic among Chabu adolescents of southwestern Ethiopia, aforager-farming society. Research with the Aka foragers questioned several evolutionary predictions about
who should innovate and how innovations should be transmitted (i.e. adolescents, males in particular,
should be more innovative than children and adults; innovations should spread by horizontal transmission
and; adolescents should pay particular attention to prestigious, “successful” peers). In-depth and structured
interviews, informal observations and systematic ranking and sorting techniques with fourteen Chabu
adolescents and eight adults were utilized in this study. Results indicate that as with the Aka data: 1) cultural
terms specific to innovation existed; 2) innovations and innovators were easily identified by adolescents; 3)innovators actively sought out individuals who exhibited pro-social qualities and were “good teachers” and; 4)
adults were more often identified by the adolescents to be innovators. One result was inconsistent with the
Aka study: Chabu adult females were more likely to be identified as innovators than were adult males.

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Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

C og niti on and c u ltur a l ni che constr uc ti on among
Neanderthals and Modern Humans
Barry S. Hewlett

Washington State University, Vancouver, Canada. Email: hewlett@vancouver.wsu.edu

This paper summarizes recent research on social learning in contemporary hunter-gatherers and how thesedata may provide insights into why Modern Humans replaced Neanderthals. First, the question as to whether
or not two cognitive mechanisms hypothesized to be essential to human cultural evolution—teaching andoverimitation—exist among contemporary foragers is considered. Second, an overview of the culturally
constructed niches of social learning in contemporary foragers is provided. The interactions between the cognitive
mechanisms and the culturally constructed niches are hypothesized to contribute to rapid, high fidelity, andinnovative social learning. Finally, comparisons between the culturally constructed niches of Neanderthals andModern Humans are utilized to explain Neanderthal replacement by Modern Humans.

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Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

Theoretical and ethnographic studies on play and learning
behaviors: The case of hunter-gatherer children in the forest
Nobutaka Kamei

Aichi Prefectural University, Japan. Email: nobutaka.kamei@gmail.com

Play and brain science
Play activities are phenomena that are widely observed both among humans and animals. It is possible toregard them as behaviors acquired in the process of evolution. It is also important to analyze and understandplay behaviors in the aspect of brain science. Recent brain science argues the relationships between categoriesof play behaviors and particular parts of the brain (for example, the positive correlation between the frequency
of social play and the size of the cerebellum among several species of mammals). Also, it is pointed out thatsome neurotransmitters and receptors are related with particular categories of play (for example, the relationshipbetween the μ-opioid receptor and the social play) (Nakagawa 2012).
The studies on play conducted by sociologists and cultural anthropologists have developed the theory of
universality and cultural particularity through the analyses of diverse play of humans (Caillois 1957; Kamei ed.
2009). This theory can be reinforced by the brain science on play. Though there exist researches that consider
play as any behaviors that are marked by meta-communication of “this is play” (Bateson 1972), these kinds ofphenomena also can be examined in the viewpoint of brain science.
Culture and play
Play is often regarded as one of subcategories of human culture in general. However, it can be considered as moreessential factors for culture. I pointed out three essential and universal aspects of play referring to classical studies
on it: (1) play is free; (2) play is done in limited time and space; (3) play can be carried out without any benefits.
If we accept the classical definition of “culture” by Edward B. Tylor (1871): “culture is … complex whole which
includes knowledge, belief, art, law, morals, custom, and any other capabilities and habits acquired by man as amember of society,” we can find two fundamental functions of play for the existence of culture.
One of the functions is for the transmission of culture. As Sperber (1996) pointed out, cultural forms are stable
and widely distributed just because children find them easy to think and easy to learn. If some activities canachieve playfulness for the next generation, such as children, these activities can be easily transmitted to them andsustained without special interferes such as education.
Another is the function for the innovation of culture. Play is always carried out in free and open rules by players.
This aspect allows them to change the behaviors anytime and any ways as they like. Everything kept andcontinued by the adults can be easily thrown away and any traditional patterns can be revolutionized by the newgeneration simply because these do not attract the youths (Platon 360-350 BC; Benjamin 1969).

090

Caillois (1957) pointed out four essential elements of play, agôn, alea mimicry and ilinx, in his theory of universal
grammar of play. These also can be analyzed and evaluated in the universal system of cultural transmission andinnovation (Kamei 2010). The human activities related to cultural transmission and innovation, such as theeducation, teaching and leaning, have to be examined by this viewpoint of play theory.

Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

Fieldwork in hunting-gathering societies
It has been pointed that hunting-gathering societies have neither educational institutions nor educational
behaviors among their children. Hunter-gatherer children have their own culture, which is connected to thetransmission of hunting-gathering culture. These were the results of my anthropological research in 1996-1998
among Baka children in the tropical rain forests in the East Region of the Republic of Cameroon (Kamei 2010).
Fifteen years later, with the further spread of schooling and the introduction of restrictions on hunting, howhave children's culture and life changed? Are the characteristics of cultural transmission in foraging societies still
maintained? This research aims to collect ethnographic data on children’s daily activities including play, hunting,gathering, fishing, housework and so on. It also aims to describe the system of cultural transmission through thisage group as one of the characteristics of foraging societies.
[Period] March 2012 (5 days); August-September 2012 (14 days)[Field site] Settlements of Baka hunter-gatherers in the East Region, Republic of Cameroon[Methods] Participant observation and interviews with Baka children. Experimental methods on playing andlearning activities were also adopted.
Results
Through participant observation and interviews among Baka children, the following points were found.
(1) School attendance rates have increased among both boys and girls, and consciousness of schooling is alreadyshared among the majority of parents.
(2) Play and subsistence activities (hunting, gathering and fishing) conducted by boys and girls observed out ofschool have not changed [Case 1].
(3) Social and cultural changes among adults partly affect children's activities, but do not affect their way ofplaying and learning [Case 2, 3].
[Case 1] Fishing by girls. They form children's groups and go into the forests. They have experiences to hunt,gather and fish in the natural environments.
[Case 2] Slide play by boys. No teaching was observed among them. The elder boys only let the younger boys
imitate their play behaviors.
[Case 3] Board game “Songo" by boys and girls. They learn how to behave first through the imitation, and how toplay the game later without teaching.
Discussion
The results show that in the past fifteen years, children's culture and life have not changed. Observation andimitation are still the essential ways of learning among children. As before, it is rare to observe positive teachingactivities among them. The characteristics of foraging societies “without education” are still maintained. Together
with these observations, a “learning model of hunter-gatherer children supported by their play sense” can bepresented in order to discuss the nature of "educational activities” by human beings.
“Play sense,” perhaps universally shared among human beings as one of the functions of the brain, has tobe examined to clarify the system of human culture. If we expect play sense as one of the factors of culturaltransmission and innovation among the humans, especially as the factor of the different history of Homoneanderthalensis and Homo sapiens, “brain science on play,” as well as “ethnographic studies on play,” will become
indispensable area of research for the investigation of human evolution.

091

Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

References
Bateson G (1972) Steps to an ecology of mind: Collected essays in anthropology, psychiatry, evolution, and epistemology. Chandler Pub. Co., San Francisco.
Benjamin W (1969) Uber Kinder, Jugend und Erziehung. Frankfurt/M.
Caillois R (1957) Les jeux et les hommes. Gallimard, Paris.
Hirschfeld LA (2002) Why don't anthropologists like children? American Anthropologist 104(2): 611-627.
Huizinga J (1938=1955) Homo ludens: A study of the play-element in culture. Beacon Press, Boston.
Kamei N (2010) Little “hunters” in the forest: Ethnography of hunter-gatherer children. Kyoto University Press, Kyoto.
Kamei N (ed) (2009) Introduction to the anthropology of play: Encounters with “children” in the fields. Showado, Kyoto.
Nakagawa A (2012) Brain and play. In: Iwata M, Kawamura M (eds) Brain and arts. Igaku Shoin, Tokyo, pp. 197-210.
Platon (360-350 BC (?)) Laws. Greece.
Sperber D (1996) Explaining culture: A naturalistic approach. Blackwell, Oxford.
Tylor EB (1871) Primitive culture. J. Murray, London.

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Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

Why do wild bonobos not use tools like chimpanzees do?
Examination of factors influencing the tool behaviors
Takeshi Furuichi1*, Crickette Sanz2,3, Kathelijne Koops4, Tetsuya Sakamaki1,
Heungjin Ryu1, Nahoko Tokuyama1 and David Morgan3,5
1. Primate Research Institute, Kyoto University, Japan; 2. Department of Anthropology, Washington University, USA; 3. Congo Program,
Wildlife Conservation Society, Republic of Congo; 4. Department of Archaeology and Anthropology, University of Cambridge, UK; 5.
Lester E. Fisher Center for the Study and Conservation of Apes, Lincoln Park Zoo, USA. *Email: furuichi@pri.kyoto-u.ac.jp

One of the most conspicuous behavioral differences among great apes is the paucity of tool use among wildbonobos (Pan paniscus) (Kano 1982; Ingmanson 1996; Hohmann and Fruth 2003) in comparison to chimpanzees(Pan troglodytes) who are one of the most prolific and skilled tool users in the animal kingdom (Sanz and Morgan
2007; Shumaker et al. 2011). This is in spite of the fact that bonobo tool use repertories are as large and diverse
as chimpanzees in captive settings (Herrmann et al. 2010; Gruber et al. 2010), suggesting that the differencesin cognitive abilities cannot fully explain the differences in tool using behaviors between these two species. This
study on the differences in tool use behaviors and related factors between closely-related ape species mightprovide insights for understanding the differences in behaviors by Neanderthals and Modern Humans.
In this study, we compared tool using behaviors and potential drivers of these behaviors in the Wamba bonobo
population located in central Democratic Republic of Congo with the Goualougo chimpanzee populationof northern Republic of Congo. The tool use repertoire of wild bonobos was comprised of only 13 behaviors,
compared to 42 for chimpanzees. However, the number of tool behaviors observed in each study site was similar
between bonobos and chimpanzees, and many types of tool use for social, self-grooming/stimulation, andcomfort/protection functions were commonly used by both species. A marked difference is that 25 of 42 tool
behaviors exhibited by chimpanzees are performed for feeding, in contrast to a single report of bonobos using aleaf sponge to drink water.
We examined whether the differences in tool use repertoires can be explained by the necessity, opportunity,
relative profitability, or invention hypotheses (Fox et al. 1999; Spagnoletti et al. 2012; Koops et al. 2013; Sanzand Morgan 2013). We found that habitat composition and fluctuation of fruit production at these two sites
were similar, particularly when compared with variation observed between sites within each species. Thus it was
unlikely that the necessity hypothesis explains the lack of tool use for feeding in bonobos. Though further study at
Wamba is needed, we did not identify any obvious differences in prey availability that would indicate differencesin tool using opportunities between the sites. This study could not test the relative profitability hypothesis, andfurther research is needed on whether tool use is the most efficient means of calorie or protein intake for wild
apes. Bonobos at Wamba formed much larger and stable parties than chimpanzees at Goualougo, which wascontrary to the prediction by the invention hypothesis.
Another explanation is that differences in tool use behavior between bonobos and chimpanzees might not be
explained by the current ecological or social conditions, but rather by circumstances during the Pleistocene
Epoch: the ability for tool use evolved only in chimpanzees under certain conditions in the past, and therefore
chimpanzees currently living in all types of habitats inherently use tools for feeding while bonobos in any type of
habitat do not. However, this hypothesis may not explain why there is no substantial between-species differencein ability for tool use under experimental conditions. The observed species differences might also reflect divergent
behavioral predispositions, rather than actual differences in cognitive abilities. Chimpanzee tool use does notseem to be the most effective means of increasing energy intake, but may be performed when individuals haveleisure time (Nishie 2011), while bonobos spend much of their leisure time in play and play is common among

093

Learning Strategies and Cultural Evolution – Anthropological and Primatological Approachesadults (Enomoto 1990; Palagi 2006). Such differences in behavioral preference may explain why bonobos arelikely to perform like chimpanzees in experimental conditions, but do not regularly perform tool use in natural
settings.
This paper is in press in Behaviour with a title “why do wild bonobos not use tools like chimpanzees do?”References
Enomoto T (1990) Social play and sexual-behavior of the bonobo (Pan paniscus) with special reference to flexibility. Primates 31:
469-480.
Fox EA, Sitompul AF, van Schaik CP (1999) Intelligent tool use in wild Sumatran orangutans. In: Parker ST, Mitchell RW, Miles
HL (eds) The mentalities of gorillas and orangutans. Cambridge University Press, Cambridge, pp. 99-116.
Gruber T, Clay Z, Zuberbuhler K (2010) A comparison of bonobo and chimpanzee tool use: Evidence for a female bias in the Pan
lineage. Animal Behaviour 80: 1023-1033.
Hohmann G, Fruth B (2003) Culture in bonobos? Between-species and within-species variation in behavior. Current Anthropology
44: 563-571.
Ingmanson EJ (1996) Tool-using behaviors in wild Pan paniscus: Social and ecological considerations. In: Russon AE, Bard KA,
Parker ST (eds) Reaching into thought: The minds of the great apes. Cambridge University Press, Cambridge, pp. 190-210.
Herrmann E, Wobber V, Call J (2008) Great apes’ (Pan troglodytes, Pan paniscus, Gorilla gorilla, Pongo pygmaeus) understanding of tool
functional properties after limited experience. Journal of Comparable Psychology 122: 220-230.
Kano T (1982) The use of leafy twigs for rain cover by the pygmy chimpanzees of Wamba. Primates 23: 453-457.
Koops K, McGrew WC, Matsuzawa T (2013) Ecology of culture: Do environmental factors influence foraging tool use in wild chimpanzees, Pan troglodytes verus? Animal Behaviour 85: 175-185.
Nishie H (2011) Natural history of Camponotus ant-fishing by the M group chimpanzees at the Mahale Mountains National Park,
Tanzania. Primates 52: 329-342.
Palagi E (2006) Social play in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): Implications for natural social systems andinterindividual relationships. American Journal of Physical Anthropology 129: 418-426.
Sanz CM, Morgan DB (2007) Chimpanzee tool technology in the Goualougo Triangle, Republic of Congo. Journal of Human Evolution 52: 420-433.
Sanz CM, Morgan DB (2013) Ecological and social correlates of chimpanzee tool use. Philosophical Transactions of the Royal Society B 368: 20120416.
Shumaker RW, Walkup KR, Beck BB (2011) Animal tool behavior: The use and manufacture of tools by animals. Johns Hopkins
University Press, Baltimore, Maryland.
Spagnoletti N, Visalberghi E, Ottoni E, Izar P, Fragaszy D (2012) Stone tool use in wild bearded capuchin monkeys, Cebus libidinosus: Is it a strategy to overcome food scarcity? Animal Behaviour 83: 1285-1294.

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Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

Some differences between humans and other animals

Magnus Enquist1*, Stefano Ghirlanda1,2 and Johan Lind1

1. Centre for the Study of Cultural Evolution, Stockholm University, Sweden; 2. Department of Psychology, Brooklyn College, New York,USA. *Email: magnus.enquist@intercult.su.se

Knowledge about the behavioural and mental abilities of animals is important for understanding human
evolution. However, it has been difficult to reach any scientific consensus about such abilities in birds or primates.
This will be illustrated with some examples including short term memory, social learning and sequential
behaviour. The second part of the talk is more theoretical and will discuss the behavioural and mental flexibilityof animals, why animals sometimes appear smarter in the wild than in the laboratory, and whether or not animals
can think. The presentation will end with some arguments why human like mental abilities have only evolved
once.

095
Learning Strategies and Cultural Evolution – Anthropological and Primatological Approaches

Population change and cultural variation

Laurel Fogarty1, Kenichi Aoki2, Joe Yuichiro Wakano2 and Marcus Feldman1*
1. Department of Anthropology, Stanford University, USA; 2. Meiji University, Japan. *Email: mfeldman@stanford.edu

The human population has undergone explosive population growth since the beginning of the Holocene
around 10,000 years ago. The effects of this population increase have been described in recent genetic studies(Keinan and Clark 2012), where it was shown that recent population expansions resulted in an increase overtime in the number of rare alleles in the population. However, little work has directly addressed the effect of
population expansion (or contraction) on the frequency distribution of cultural traits in the population. Here
we introduce a cultural ‘trait frequency spectrum’ in direct analogy to the genetic ‘site frequency spectrum’ andexamine the effects of population growth or decline on the distribution of cultural trait frequencies in a population. We use the simulation framework outlined by Fogarty et al. (in press) and compare the trait frequency
spectra for populations using random oblique cultural transmission and best-of-k cultural transmission, elucidating the effects of population size as well as population growth. Finally, we comment on the implications ofthese findings for our understanding of human cultural evolution and cultural accumulation over time.

096
Learning Strategies and Cultural Evolution – Perspectives from Theoretical ModellingWhy did the arts precede prehistoric settlement in Modern
Humans ?
Shiro Horiuchi

Yamagata University, Japan. Email: horiuchi.shiro@gmail.com

Introduction
Why did only our species, Homo sapiens, leave behind artistic products, whereas other homo species, including
Neanderthals, did not? The producers of art, or artists, might have suffered from much opportunity cost.
One hypothesis is that different groups might have communicated well because of artists, who functioned ascoordinators who connected individuals of different groups (Conkey 1980). Through the role that artists played,other groups might have been able to settle in an area and develop their own cultural traits. I introduce an agentbased model (ABM) simulation to test cultural accumulation in the time of the Replacement (Horiuchi andKubota 2013), in order to simulate the dynamics of arts and settlement.
Methods
The ABM assumes 100 agents. They are uniformly distributed on the 10 × 10 cells. There are 10 × 10 cultural
traits, ci, each of which is unique to each cell, and is cumulative. Each agent gains cultural traits as the 10 × 10
matrix, C (ci ∈C).
The agents have 10 units of effort. Each agent allocates his efforts among Hunter-gather (H), Settlement (S),and Arts (A). If an agent allocates eh efforts to H, he gains the cultural trait of his residential cell, as well as thoseof his surrounding cells. Now, we define the distance between cell i and cell j as xi,j. The agent of cell i gains thexcultural trait of j as eh p i,j , where p is the discounting ratio of H (0 < p ≤ 1). If he allocates es efforts to S, he gains
the cultural traits of his residential cell as esq, where q is the effect of settlement (q ≥ 1). If an agent allocates ea
efforts to A, he may be able to communicate with other agents from surrounding areas. Now the agent i allocatesea,i efforts to A. He communicates with the agent j if the value ea,i + ea,j – rxi,j > 0 and is maximum, where r is thecost of communication (r > 0). While communicating, the two agents learn cultural traits from each other by thediscounting ratio s; i.e., we assume that the cultural trait k of the agents i and j is ck,i and ck,j, respectively. If ck,i < sck,jor sck,i > ck,j, the cultural trait k of the agent i or j is improved up to sck,j or sck,i, respectively (0 < s < 1). Naturally eh +x
es + ea = 10. The payoff of the agent is equivalent toΣcj u i,j , where u is the discounting ratio of other cultural traits (0< u < 1).
At the initial condition, all agents allocate all their efforts to H. In the simulation, the agents change their
allocation of efforts successively through trial and error, to maximize their payoff. The ABM verifies the conditionand process by which the agents allocate their efforts to S or A.
Results
Since the ABM has many independent variables, p, q, r, s, and u, herein I set the parameters (q, r, s, u) = (2, 1, 0.5,0.5) and change only p to simplify the analysis. When p = 1, all agents keep their efforts at (H, S, A) as (10, 0, 0).
When p = 0.5, however, an increasing number of agents allocates their efforts from H to S as time passes, whilea few agents allocate their efforts from H to A (Fig. 1). A communication network appears where artists, who097

Learning Strategies and Cultural Evolution – Perspectives from Theoretical Modellingallocate much effort to A, are situated at the hub positions (Fig. 2). At t = 30,000, the agents are classified into
three groups on the basis of their position in the network (Fig. 2b); the agents of each of the three groups haveunique sets of cultural traits that are distinguishable from other groups.
I ran 20 simulations for each p, from 1 to 0.2, and verified the results at t = 30,000. Figs. 3, 4, and 5 respectively
show the box plot of the payoffs of agents, the ratios of agents who belong to the largest group, and the sum of
the cultural trait of each cell gained by the residential agent.
Discussion
If agents are less likely to gain cultural traits by H, they start to shift their efforts from H to S or A. Many agents
allocate all their efforts to S, whereas a few agents specialize in A. Although they gain few benefits from H, they
may gain much payoff by shifting their efforts to S and are coordinated by the agents of A. The agents of A work
well as coordinators, who let other agents allocate their efforts to S.
The present ABM suggests the process of civilization. First, some individuals work as artists and coordinate
communication among different groups; then, most individuals start a settlement and more cultural traits
develops in the entire population. The reason why individuals started creating art is that there were few benefits
from hunting-gathering because of severe environmental conditions, such as during the glacial period. In thepresentation, I show how the agents of A are distributed through computer simulations. Future research may
predict the areas where artistic products were made by Modern Humans in prehistoric times.

Fig. 1 Average efforts to H, S and A of
all agents along time (0 < t < 100,000).

Fig. 2 Communication network coordinated byartists. The hub agents really allocate much efforts
to A. (a) t = 10,000. (b) t = 30,000.

Fig. 3 Average payoffs of agents at
different p.

Fig. 4 Maximum cluster of agents at
different p.

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Learning Strategies and Cultural Evolution – Perspectives from Theoretical ModellingFig. 5 Summed cultural traits by agentsat different p.

Population and culture: A theoretical basis for dramatic
regime shifts?
Kenichi Aoki

OSCRIP, Meiji University, Japan. Email: kenaoki@meiji.ac.jp

Model and analysis
Malthus proposed that “population equilibrates with resources at some level mediated by technology,” whereas
Boserup believed that “technological change is itself spurred by increases in population” (quoted from Lee 1986).
Many Palaeolithic archaeologists currently emphasize the Boserupian premise in interpreting major changes instone tools or other cultural artefacts. In the seminal paper cited above, Lee presents a coevolutionary model for
population and technology that synthesizes the contrasting viewpoints of Malthus and Boserup. Richerson andBoyd (2013) have recently discussed the importance of adopting such an approach in understanding progressive
and regressive cultural changes during the Palaeolithic. Here, we describe and analyze a minimal mathematicalmodel, in the spirit of Ghirlanda and Enquist (2007), that instantiates the verbal model of Richerson and Boyd(2013).
For a population let us denote its mean cultural (technological) level by and its size
We assume the dynamic> 0 and its size by N > 0.

(1)
for the mean cultural level resulting from error-prone social learning, whereandare positive parameters,and ε≈ 0.577 is Euler’s constant (Henrich 2004; Mesoudi 2011). Next, assume the logistic model of population
growth,(2)

where, K , and D are positive parameters. The assumption here is that the cultural level of an individual
does not contribute to the biological fitness of that individual, perhaps because resources are shared, but thatthe mean cultural level of a population, , determines—has a positive effect on—the carrying capacity of the
population.
Make the transformation of variable, which yields

099

Learning Strategies and Cultural Evolution – Perspectives from Theoretical Modelling

(3)
and
(4)

The equilibria of the system, which must simultaneously satisfy
intersection(s) of the exponential curve, are given by the
(5)

where

, with the sigmoid curve(6)

Fig. 1 below depicts the phase plane for the model with the transformed cultural level,, on thehorizontal axis and population, , on the vertical axis. The parameter values are λ=0.2, c =6, z*=18, K=10, D=30.
In this case, the blue curve which is the plot of Eq. (5) (the Boserup line) and the red curve which is the plot of
Eq. (6) (the Malthus line) intersect three times to yield three valid equilibria (that satisfy z > 0 ).
The directions of change in the various regions of the phase plane are given by Eq. (3) and Eq. (4) and indicatedby black arrows. They show that the two outside equilibria—the ones on the right and left correspond to highand low culture regimes, respectively—are locally stable whereas the middle equilibrium is unstable. Hence, wehave bistability in this case.
Fig. 2 below withon the horizontal axis andon the vertical axis shows that this numericalexample exhibits hysteresis. Parameter is a measure of innovativeness, with smaller values corresponding to greater
innovativeness (Kobayashi and Aoki 2012). The other parameters are set atThe
two solid lines denote alternative locally stable equilibria—high and low culture regimes—whereas the broken
line gives the unstable equilibria.
Fig. 2 suggests the following scenario for changes in cultural level when an evolutionary increase in innovativenessoccurs ( decreases). Initially, there will be a gradual increase in cultural level along the lower solid line
representing equilibria at the lower cultural level. Then, when
passes a critical thresholdknown as a“catastrophic bifurcation”, there will be a regime shift to an equilibrium at the higher cultural level represented by
the upper solid line. Empirically, this sudden shift may be observable in the archaeological record as a “creative
explosion.” If for some reason innovativeness subsequently declines, the cultural level will gradually decrease
along the upper solid line of equilibria. Then, when
passes the other critical threshold, a regime shift
occurs to the lower stable line. The forward and backward switches follow different paths, i.e. we have hysteresis.

100

Hysteresis is also observed when the “cultureless” carrying capacity,
constant.

Learning Strategies and Cultural Evolution – Perspectives from Theoretical Modelling, is varied with the other parameters held

References
Ghirlanda S, Enquist M (2007) Cumulative culture and explosive demographic transitions. Quality and Quantity 41: 591-600.
Henrich J (2004) Demography and cultural evolution: How adaptive cultural processes can produce maladaptive lossess—The
Tasmanian case. American Antiquity 69: 197-214.
Kobayashi Y, Aoki K (2012) Innovativeness, population size and cumulative cultural evolution. Theoretical Population Biology
82: 38-47.
Lee R (1986) Malthus and Boserup: A dynamic synthesis. In: Coleman D, Schofield R (eds) The state of Population Theory.
Basil Blackwell, pp. 967-130.
Mesoudi A (2011) Variable cultural acquisition costs constrain cumulative cultural evolution. PloS One 6: 1-10.
Richerson PJ, Boyd R (2013) Rethinking paleoanthropology: A world queerer than we supposed. In: Hatfield G, Pittman H (eds)
Evolution of mind, brain and culture. University of Pennsylvania Press, Philadelphia, pp. 263-302.

Fig. 1

Fig. 2

101
Learning Strategies and Cultural Evolution – Perspectives from Theoretical ModellingLearning schedule and cultural evolution: COS and ESS

Joe Yuichiro Wakano

Meiji University, Japan. Email: joe@meiji.ac.jp

Introduction
In the evolution of Modern Humans, innovative stone tools might have played an important role. Advanced andcomplex lithic industries can never be invented by a single individual without building on knowledge acquired
from others. They are the results of cumulative cultural evolution in which knowledge is inherited from parental
generations and passed on to descendant generations. However, if all individuals simply copied what was alreadyknown, there would be no advancement in technology. Contribution to culture by improving on the precedingknowledge, such as a discovery of new adaptive use or form of stone tools, is also crucial for cultural evolution.
Thus, it is an important problem to determine what learning strategy can support the cumulative culture thatdevelops the fastest: when and how should an individual perform learning during one's life?
The balance between the time allocated to learning and the time allocated to the exploitation of the learned
knowledge is then the next question. To maximize the accumulation rate of culture, one should spend sufficient
time to absorb the existing knowledge in the parental generation and then spend all the rest of the lifetime toimprove the knowledge by individual learning. However, such a learning schedule does not necessarily maximize
the fitness of an individual, i.e., the expected number of biological offspring. The ultimate goal for an individual is
the maximization of the benefit provided from the knowledge or culture, not the maximization of the knowledge
itself.
Previous theoretical studies
Theoretical researches surveying the evolution of leaning as life-history strategies have started only recently.
One of the well-known studies on a leaning schedule is Social Learning Tournament (Rendell et al. 2010).
However, the Tournament included so many factors and the analytic treatment is impossible. Enquist et al. (2007)
compared the performances of pure SL strategy, pure IL strategy, and “critical social learner” who performs
individual learning (IL) only if social learning (SL) had failed to achieve OK solution. They showed that criticalsocial learner outperforms the pure strategies and thus evolves. Aoki et al. (2012) performed a more exhaustive
study of a two-stage model in which any mixture of SL and IL is allowed in each stage and the environment
may fluctuate. They showed that in a constant environment pure SL followed by pure IL is evolutionarily stablestrategy when the efficiency of SL is not too low.

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In a series of studies on the evolutionarily stable learning schedule, Lehmann et al. (2013) studied a casewhen trade-off between learning and exploiting exists. This model is complex as it includes many factors (i.e.,
continuous time axis, horizontal transmission, and environmental changes). One of their results is that, whenenvironmental change is negligibly rare, the equilibrium of the cultural level realized by the evolutionarily stablelearning schedule is not very high compared to the level realized by the pure IL strategy. This result is counterintuitive because improvement achieved by IL should accumulate over generations when environmental change
is rare (a sufficiently small decay rate of information), and thus we naturally expect the cultural level to increase
to reach the maximum level that is transmittable to the next generation (limited only by the efficiency of SL).

Learning Strategies and Cultural Evolution – Perspectives from Theoretical Modelling

However, the predicted cultural level was far below this level.
Aim of study
The goal of the present study is to clarify the factors that facilitate or suppress the cumulative cultural evolution
over generations. When and how does the maximization of individual fitness (natural selection) also maximize
the cultural level of the society after a sufficient number of generations? For this purpose, we study a relatively
simple model with a two-stage life history when there is trade-off between learning and exploiting.
Model
A learning schedule considered here is a broad developmental pattern over a single individual's lifetime in the
differential use of IL and SL with regard to behaviors that may take a long time to be acquired. It differs from theordered application of IL and SL in skill acquisition at any one time, which may occur repeatedly during one's
lifetime.
Allocation of effort is the evolving strategy in this study. In the first stage, an individual performs IL with effort
and SL with effort 1-u0. In the second stage, the individual allocates the learning effort to IL and SL with a ratio
u1:1-u1. During this second stage, the individual can reproduce offspring. Reproductive success is determined by
two factors; cultural level and the effort put in reproduction. Most animals including human can learn something
before the body sexually matures. It also seems adaptive to allocate more learning effort in the earlier stage of life
than allocating learning effort equally in all life stages. Thus, we assume that an individual does not reproduce
in the first stage. In the second stage, the fraction v of time is dedicated to learning, and the fraction 1-v toexploiting the knowledge for reproduction. Thus in the second stage, efforts u1v, (1-u1)v, and 1-v are allocated toIL, SL, and exploitation, respectively. The life history strategy is represented by the triplet (u0,u1,v).
ResultWe are interested in a case when the efficiency of SL is sufficiently large. Otherwise cultural accumulation neveroccurs. If there is no trade-off between learning and exploitation, a learning schedule performing SL then IL is
predicted. Thus, if the SL efficiency is sufficiently high, the equilibrium of the cultural level can increase to almost
infinity. However, when a trade-off between learning and exploitation exists, COS and ESS are completely
different.
Coordinated-Optimal Strategy (COS)
COS is such a learning strategy that maximizes the equilibrium value of fitness when all individuals in population
adopts this strategy. The idea is similar to the concept of the Pareto-optimality where payoff of a certain player
cannot increase without decreasing payoff of another player. COS would be realized when all individuals behave
as if they would cooperate to maximize the fitness (benefit) of a group in far future. In a simple model (Wakano
and Miura 2014), COS predicts cultural accumulation.
Evolutionarily Stable Strategy (ESS)
ESS is such a learning strategy that any other strategy yields lower fitness when all individuals adopts thisstrategy. It is sometimes referred to as undefeatable strategy. ESS would be realized when all individuals behave
as if they would try to maximize the fitness (benefit) of himself/herself (and hence, interested only in immediate
benefit, not future). In a simple model (Wakano and Miura 2014), ESS predicts no cultural accumulation.

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ELearning Strategies and Cultural Evolution – Perspectives from Theoretical Modelling

Discussion
If learning schedule is genetically determined, the ESS should evolve. In a simple model, it is IL-only strategy.
Thus, cumulative culture never evolves. The model neglects so many factors such as cooperative informationtransfer among relatives (e.g., from parent to offspring), thus such factors might explain the observed cumulative
culture in Modern Humans. Another possibility is that learning schedule itself is cultural trait, i.e. learned. Under
this hypothesis, the COS might be predicted, but it will result in a highly hierarchical model which is difficult toanalyze. Human developmental pattern (e.g., body and brain growth curve) represents the restriction on learning
behavior of an infant/child/juvenile and it must have close relationship with human learning schedule. The
connection between the study on learning schedule and the study of human development might provide a way to
explain the evolution of cumulative culture.
References
Enquist M, Eriksson K, Ghirlanda S (2007) Critical social learning: A solution to Roger’s paradox of nonadaptive culture.
American Anthropology 109: 727-734.
Rendell L, Boyd R, Cownden D, Enquist M, Eriksson K, Feldman MW, Fogarty L, Ghirlanda S, Lillicrap T, Laland KN (2010)
Why copy others? Insights from the social learning strategies tournament. Science 328: 208-213.
Aoki K, Wakano JY, Lehmann L (2012) Evolutionarily stable learning schedules and cumulative culture in discrete generation
models. Theoretical Population Biology 81: 300-309.
Lehmann L, Wakano JY, Aoki K (2013) On optimal learning schedules and the marginal value of cumulative cultural evolution.
Evolution 67: 1435-1445.
Wakano JY, Miura C (2014) Trade-off between learning and exploitation: The pareto-optimal versus evolutionarily stable learning schedule in cumulative cultural evolution. Theoretical Population Biology 91: 37-43.

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Learning Strategies and Cultural Evolution – Perspectives from Theoretical Modelling

DAY 6: December 5, 2014

Cognition and Learning
Looking into the Neanderthal Brain
Closing Remarks

The beginning of symbolic art and the learning cycle innatureHideaki Terashima

Kobe Gakuin University, Japan. Email: hideaki_terashima@icloud.com

A bunch of modern behavior that characterize Modern Humans has developed gradually in the Middle Stone Ageof Africa after the emergence of “anatomically Modern Humans” around 200 ka (McBrearty and Brooks 2000).
There were various objects supposed to have symbolic meaning such as burial accompanied with artifacts, ornaments
like beads and pendants, cave art and fine sculptures, and so on. For beads and pendants, the date of fabrication
seems to go back to 40-50 ka, and some ostrich eggshell beads were found as fabrication of 83 ka. Pigments seems
to have been in use from more than 50 ka. On the other hand, artistic paintings and sculptures had not appeared
until the upper paleolithic in Europe.
The production of symbolic art has diffused in Europe rapidly after the replacement of Neanderthals by Modern
Humans. The oldest cave paintings were found at Chauvet in southwestern France, which is dated back to ca.
32,000 BP. The most conspicuous feature of early cave art is that the paintings mostly consisted of large animals
such as bisons, aurochs, horses, red-deers, reindeers, and ibexes being depicted in an astonishingly vivid and beautiful
forms. Human pictures are extremely rare and they are mostly represented in a childish manner, seriously abstracted
and deformed, and sometimes as a therianthropic appearance. The reason why people began to create such artistic
representation after the replacement is not clear at all. But judging from the accurate and vivid animal images, it
seems doubtless that people who created such paintings had great interest to nature, in particular to large animals
living in their vicinity. It seems that humans have awaken to the splendidness of nature and animals. Before thereplacement period animals were just targets of hunting for humans, either modern or Neanderthal, and sometimearound the replacement, utterly different ideas about animals were born and developed in human mind.
Today’s hunter-gatherers have two approaches to wild animal world: one is natural historical approach and the otheris sympathetic approach. These two approaches make up the most important part of the life of hunter-gatherers.
Hunter-gatherers go stroll about nature and explore the environment for new information everyday: people pickup
any changes in the environment such as the new traces of animal footprints, fruit ripening smell, color of tree
blossoms, the sound and movement of honeybees, and so on. As getting various information from the environment
people make prediction following indigenous eco-models or theories about natural phenomena and adjust theirforaging plan. If the prediction comes true, the information is recognized to be useful and accumulated in theirknowledge archives. The accuracy and wideness of knowledge about nature in hunter-gatherers are well documented
by many ethnologists and biologists: one report says that “the !Kung appear to know a good deal more about many
subjects than do the scientists.” (Blurton-Jones and Konner 1976).
On the other hand, hunter-gatherers have a quite different approach, a sympathetic one to animals. Although
animals are the objects of hunting activity, they are also admirable companions sharing the same natural environment
with humans and showing abilities which surpass humans. People try to understand the animals sympatheticallythrough intuition, mind-reading or theory of mind, and anthropomorphic interaction. Among hunter-gatherers
it is quite commonly believed that humans can communicate with animals and that makes a key point in hunting
process.

106

The sympathetic approach to animals opens a new way to human learning behavior. Pedagogy is one of the useful

Cognition and Learningmethods of learning among humans, which consists of interaction between the teacher and the learner in an
educational context. The relationship between the teacher and the learner is crucial point for obtaining expected
results in pedagogy. Outside the school institution, the teacher is not confined to experts, and even those who
simply watch over the learner can be a teacher as long as there is mutual recognition between the teacher and thelearner. And this pattern can be extended even to animals. For example, a hunter who wants to understand the
behavior of bears can learn from the bear by regarding the bear as the teacher, following it and imitating everything
it does. In repeating such learning, the hunter will finally find himself totally behaving like the bear. The hunter
becomes to know what the bear will do next moment. This is a kind of imitative social learning and also a kind ofmeta-learning in which the learner has to monitor ones own learning process all the time following and imitating
the animal.
Either by the natural historical approach or the sympathetic one, hunters place themselves deep in the environmentand hear sounds and feel moves of nature. The two approaches look like two different ways to arrive totally different
goals, but actually they interrelate with and complement each other to make an organized body of knowledge called
TEK (traditional ecological knowledge.) It is a synthetic knowledge consisting not only of natural, but also of socialand cultural elements.
Nature being marked with its miraculous complexity, richness of detail and variability, brings humans infinite
opportunity for discovery and new learning. It may not be an exaggeration to say that all ideas of human beings,
thoughts and imaginations have been derived from nature. There is a learning cycle for hunter-gatherers in relation
to their natural environment. It starts with enormous interest in nature that motivates hunters to explore the naturalenvironment with careful attentions and observation. Acute and meticulous observation brings about discovery
which turns into knowledge and understanding. The ability of discovery gives birth to creative behavior and arouse
more interest in nature.
The intimate knowledge about nature and animals accompanied with excitement of discovery might have awaken
the upper paleolithic hunter-gatherers to create the splendid cave art. The learning cycle in nature is supported bycognitive development, theory of mind or mind reading, flexible or fluid intelligence, and meta-learning. Sociality is
another indispensable factor in that cycle because the communication with nature is based on the idea of relatedness
between humans and animals and other nonhuman creatures in nature.
References
Blurton-Jones N, Konner MJ (1976) !Kung knowledge of animal behavior. In: Lee RB, DeVore I (eds) Kalahari hunter-gatherers. Harvard University Press, Cambridge, pp. 325-348.
McBrearty S, Brooks AS (2000) The revolution that wasn’t: A new interpretation of the origin of Modern Human behavior. Jounal of
Human Evolution 39: 453–563.

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Cognition and Learning

The ability of objectification as the basis for cultural learningand creativity: Teasing as the device for training children in
Canadian Inuit society
Keiichi Omura

Graduate School of Culture and Language, Osaka University, Japan. Email: BXQ06636@nifty.com

The purpose of this presentation is to examine socio-cultural backgrounds of the Inuit adults’ teasing children
and disclose its real purpose behind it in order to consider the evolutionary basis of the learning ability of Modern Humans and thus build the hypothesis about the difference in learning ability between Neanderthals andModern Humans. Firstly, based on the Bateson’s model regarding the evolution of learning, I will examine thehypothesis of ‘cumulative cultural evolution’ proposed by Tomasello in order to modify Tomasello’s model toinclude the evolutionary process of learning ability from Neanderthals to Modern Humans. Then, I will return
to some examples of the Inuit adults’ teasing children in order to understand its characteristics. Moreover, I willsituate it in the socio-cultural backgrounds of the learning process of Inuit children and show that the teasing
functions as the device for education of the art of patience, which is the emotional basis for observation learningand creative invention. Finally, based on the results of these analyses, I will propose the hypothesis that the mostimportant ability required for full achievement of ‘cumulative cultural learning’ is the ability to objectify andmanipulate the relationships between oneself and the environment, which might distinguish Modern Humans
from Neanderthals.

108
Cognition and LearningNeural correlates of coping strategy for boredom: Its
relationship with creativity
Motoaki Sugiura1, 2*, Sugiko Hanawa1, Takayuki Nozawa3, Rui Nouchi1,2,
Ryoichi Yokoyama1, 4, Yuka Kotozaki3, Tsuyoshi Araki3 and Ryuta Kawashima1, 3
1. Department of Functional Brain Imaging, IDAC, Tohoku University, Japan; 2. Human and Social Response Research Division, IRIDeS,
Tohoku University, Japan; 3. Smart Ageing International Research Center, IDAC, Tohoku University, Japan; 4. Japan Society for the
Promotion of Science, Japan. *Email: motoaki@idac.tohoku.ac.jp

Introduction
Boredom is a negative emotion that is characterized by the lack of meaning one feels from repeated exposure to
the same stimulus or repeated performance of the same actions. Coping strategies for feeling bored are candidate
cognitive processes that may explain the creativity of Homo sapiens. Technological innovation and culturaldevelopment may be in part the result of coping against feeling bored in humans. Functional neuroimaging
studies have recently started exploration of the neural mechanism of boredom (Malkovsky et al. 2012; Mathiak
et al. 2013), while the neural bases of the coping system for boredom is yet to be addressed. In this study, we
addressed specifically the approach coping, which is to somehow cope with the stress rather than to run awayfrom it. There are two types of strategies in approach coping: behavioral coping (BC) and cognitive coping (CC)strategies (Holahan et al. 1987; Nett et al. 2010). The BC strategy involves tying to actually change the stressful
situation of boredom, while the CC strategy involves changing one’s own perception of the situation. The purposeof this study was firstly to clarify the neural mechanisms of the two strategies of approach coping with boredomusing functional Magnetic Resonance Imaging (fMRI). We also attempted to clarify the association between theneural correlates of the coping strategies and creativity.
Methods
Forty-six healthy right-handed university students participated in this study. Written informed consent was
obtained from each subject and the protocol was approved by the Ethics Committee of Tohoku University. In the
MRI scanner, the subjects alternately performed an Appreciation task, in which they viewed a visual stimulus,
such as a photo or drawing (4s), and an Evaluation task, in which the subjects rated their degree of boredom
during the preceding Appreciation task using a four-grade scaling (1: Want to see the same stimulus – 4: Want
to see a new stimulus; 1s). When the subject selected 1, 2, or 3 in the Evaluation task, the same stimulus waspresented in the subsequent Appreciation task. When the subject selected 4, in the subsequent Appreciationtask, he/she could see a new stimulus at 50% probability. When a new stimulus was not presented, the subject
selected 4 again and a new stimulus was presented in 50% of the cases in the following Appreciation task. We
hypothesized that BC activity would occur just before the first selection of 4 for each stimulus, and CC activity
would occur at the second or more presentations of the same stimuli after the first selection of 4. We analyzed
neural activity during the appreciation task on SPM8 using a GLM with the following four regressors: (1)average response to the stimuli, (2) parametric modulation on the response by the boredom rating, (3) activity
just before the first selection of 4 for each stimulus, and (4) activity for the second or later presentations of thesame stimuli after the first selection of 4. The regressors (2), (3), and (4) capture the neural response relevant to
the feeling of boredom, the BC strategy, and the CC strategy of the boredom coping, respectively. To examine
the association between the neural correlates of the coping strategies and creativity, across-subject voxelwise
single-regression analysis with the creativity score was adopted to the estimates for the regressors (3) and (4). The
creativity score was measured using the S-A creativity test (Society for Creative Minds 1969), which assesses thedegree of divergent thinking (Guilford 1967).

109

Cognition and LearningResults
The data of 45 subjects (19 females, aged 20.6±1.96) were analyzed. The neural response to the boredom feelingwas identified in the bilateral supramarginal gyrus (SMG)/angular gyrus (AG), the right superior/middle
frontal gyrus, the right precuneus, the bilateral inferior frontal gyrus, the left orbital surface (Fig. 1). Activation
during the boredom coping using the BC and CC strategies are shown in Figs. 2 and 3. Irrespective of thestrategy, neural activity related to boredom coping was identified in the ventromedial prefrontal cortex (vmPFC).
Comparing the two different strategies, neural responses specific to the BC strategy were observed in the left
post central gyrus, bilateral anterior cingulate cortex (ACC), cuneus. Those to the CC strategies were identified in
the bilateral middle temporal gyri (MTG), the right insula, the dorsomedial prefrontal cortex, the right posterior
cingulate cortex, and the right AG. As a result of regression analysis, we found a significant positive effect of
the creativity score on the estimates for the regressor (4) alone in the right superior temporal gyrus/lolandic
operculum/insula (Fig. 4).
Discussion and conclusion
We found the neural mechanism for coping with boredom, as well as the neural response related to the boredom
feeling. The results suggest that the vmPFC is related to the boredom coping irrespective of the strategies. We
consider this area to select the coping strategy: behavioral or cognitive. Previous studies suggested that thisarea is involved in the control of attention toward the external environment or the execution of ambivalentevaluations (Gilbert et al. 2006; Nohlen et al. 2013). Furthermore, the vmPFC is considered to be essential forthinking about the future, given its relevance to delay discounting (Cooper et al. 2013). The left post central
gyrus and ACC identified for the BC strategy seemed to be relevant to behavioural output considering theirknown role in the motor preparation (Zang et al. 2003). Bilateral MTG identified for the CC strategy appears
to play a key role in boredom-driven creativity based on the role of the regions in the semantic processing (Ruffet al. 2008) and execution of a creative task (Zhao et al. 2013). The observed relationship between the creativity
score and activation for the CC strategy in the right superior temporal gyrus/Rolandic operculum/insula in CC
suggest that the emotional processing implicated in this region (Craig 2002, 2011) may be a key mediator of theboredom-driven creativity. These findings support the importance of boredom and the cognitive coping strategy
for it in the creativity of Homo sapiens and suggest possible neural mechanism underlying these dynamics.

110

References
Cooper N, Kable JW, Kim BK, Zauberman G (2013) Brain activity in valuation regions while thinking about the future predicts individual discount rates. The Journal of Neuroscience 33: 13150-13156.
Craig AD (2002) How do you feel? Interoception: The sense of the physiological condition of the body. Nature Reviews Neuroscience 3(8): 655-666.
Craig AD (2011) Significance of the insula for the evolution of human awareness of feelings from the body. Annals of the New York
Academy of Sciences 1225: 72-82.
Gilbert SJ, Simons JS, Frith CD, Burgess PW (2006) Performance-related activity in medial rostral prefrontal cortex (area 10) during
low-demand tasks. Journal of Experimental Psychology-Human Perception and Performance 32: 45-58.
Guilford JP (1967) The nature of human intelligence. McGraw-Hill Companies, New York.
Holahan CK, Holahan CJ (1987) Self-efficacy, social support, and depression in aging: A longitudinal analysis. The Journal of Gerontology 42(1): 65-68.
Mathiak KA, Klasen M, Zvyagintsev M, Weber R, Mathiak K (2013) Neural networks underlying affective states in a multimodal
virtual environment: Contributions to boredom. Frontiers in Human Neuroscience 7: 820.
Malkovsky E, Merrifield C, Goldberg Y, Danckert J (2012) Exploring the relationship between boredom and sustained attention.
Experimental Brain Research 221: 59-67.
Ulrike N, Thomas GT, Lia MD (2010) What to do when feeling bored? Students' strategies for coping with boredom. Learning andIndividual Differences 20(6): 626-638.
Nohlen HU (2014) Evaluating ambivalence: Social-cognitive and affective brain regions associated with ambivalent decision-making.
Social Cognitive and Affective Neuroscience 9(7): 924-931.
Ruff I, Blumstein SE, Myers EB, Hutchison E (2008) Recruitment of anterior and posterior structures in lexical-semantic process-Cognition and Learninging: An fMRI study comparing implicit and explicit tasks. Brain and Language 105(1): 41-49.
Society for Creative Minds (1969) Manual of S-A creativity test. Tokyo Shinri Corporation, Tokyo.
Zang Y, Jia F, Weng X, Li E, Cui S, Wang Y, Hazeltine E, Ivry R (2003) Functional organization of the primary motor cortex characterized by event-related fMRI during movement preparation and execution. Neuroscience Letters 6, 337(2): 69-72.
Zhao Q, Zhou Z, Xu H, Fan W, Han L (2014) Neural pathway in the right hemisphere underlies verbal insight problem solving.
Neuroscience 256: 334-341.

Fig. 1 Neural activity related to the feeling of boredom.

Fig. 2 Neural activity during the BC.

Fig. 3 Neural activity during the CC.

Fig. 4 Correlation between activation during the CC and creativity (S-A) score.

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Cognition and Learning

The evolution of the parietal areas in Neanderthals andModern Humans: From bones to mind
Emiliano Bruner

Centro Nacional de Investigación sobre la Evolución Humana, Burgos, Spain. Email: emiliano.bruner@cenieh.es

The major form change associated with the evolution of the Modern Human brain is a marked dilation of
the upper parietal surface. An early post-natal morphogenetic stage in Homo sapiens is responsible for thisparietal bulging, and this stage is absent in chimpanzees and Neandertals. Such evolutionary change has been
hypothesized to be associated with deep parietal elements, like the precuneus or the intraparietal sulcus. Bothareas are involved in visuospatial integration. According to theories of Extended Mind, cognition is deeply
rooted into the body experience, being the body the interface between brain and environment. In this process,visuospatial integration functions are particularly important, managing the integration between inner (body)and outer (environment) coordinates. The main ports of such interface are the eyes and the hands, and theintraparietal sulcus represents a main node of their coordination. Dental evidence suggests that Neandertals usednormally to rely on the mouth for praxis, although such choice put seriously at risk its essential functions. Taking
into account that Neandertals had a complex culture but lack the parietal dilation displayed by Modern Humans,
we can wonder whether their visuospatial capacity required the involvement of an additional component in theirbody interface, because of a mismatch between biological and cultural evolution. This hypothesis in cognitivearchaeology may be tested and evaluated considering other visuospatial behaviours in extinct hominids, as well as
investigating in Modern Humans the visuospatial functions and the eye-hand system according to the principles
of extended mind.
References
Bruner E, Lozano M (2014) Extended mind and visuo-spatial integration: Three hands for the Neandertal lineage. Journal of Anthropological Science 92: 273-280.
Bruner E, de la Cuétara JM, Masters M, Amano H, Ogihara N (2014) Functional craniology and brain evolution: From paleontology
to biomedicine. Frontiers in Neuroanatomy 8: 19.
Clark A (2007) Re-inventing ourselves: the plasticity of embodiment, sensing, and mind. Journal of Medicine and Philosophy 32:
263–282.
Clark A (2008) Supersizing the mind. Emdodiment, action, and cognitive extension. Oxford University Press, Oxford.
Gunz P, Neubauer S, Maureille B, Hublin JJ (2010) Brain development after birth differs between Neanderthals and Modern Humans. Current Biology 20: R921-R922.
Iriki A, Sakura O (2008) The neuroscience of primate intellectual evolution: Natural selection and passive and intentional niche construction. Philosophical Transactions of the Royal Society B 363: 2229-2241.
Iriki A, Taoka M (2012) Triadic (ecological, neural, cognitive) niche construction: A scenario of human brain evolution extrapolating
tool use and language from the control of reaching actions. Philosophical Transactions of the Royal Society B 367: 10-23.
Malafouris L (2010) The Brain-Artefact Interface (BAI): A challenge for archaeology and cultural neuroscience. Social Cognitive
and Affective Neuroscience 5: 264-273.
Malafouris L (2010) Metaplasticity and the human becoming: Principles of neuroarchaeology. Journal of Anthropological Science
88: 49-72.
Malafouris L (2013) How things shape the mind: A theory of material engagement. MIT Press, Cambridge.
Neubauer S, Gunz P, Hublin JJ (2009) The pattern of endocranial ontogenetic shape changes in humans. Journal of Anatomy 215:
240-255.
Neubauer S, Gunz P, Hublin JJ (2010) Endocranial shape changes during growth in chimpanzees and humans: A morphometricanalysis of unique and shared aspects. Journal of Human Evolution 59: 555-566.

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Estimating the extent of brain subdivisions using skulls

Yasushi Kobayashi1*, Toshiyasu Matsui1 and Shigeru Kosuda2

1. Department of Anatomy and Neurobiology, National Defense Medical College, Japan; 2. Department of Radiology, National Defense
Medical College, Japan. *Email: yasushi@ndmc.ac.jp

Understanding the structure and cognitive functions of fossil hominid brains is one of the most important
issues in anthropology. However, due to the lack of the soft tissue in fossil skulls, we need to infer the structureand functions of the brains indirectly based on the skull morphology. To infer the outer shape of the brains,researchers have been using endocranial casts. Although meninges and cerebrospinal fluids intervene between theskull and the brain, they are relatively thin compared to the skull size, and allow approximate estimation of thevolume and the outer shape of the brain. To obtain further information from the skull, we need to infer the sulcal
and gyral patterns of the brain and determine the extent of major subdivisions of the cerebral cortex. We thus
focus on the correspondence of endocranial impressions to cerebral sulci and gyri, as well as on the relationshipbetween the locations of cerebral sulci and cranial sutures.
The sulcal and gyral patterns of the brain were clearly observed on the endocasts in prosimians and in smallsized species in anthropoids. However, there is a long history of discussion on the relationship of endocranial
impressions to cerebral sulci and gyri in hominids. Several of the earliest studies in hominid skulls showed
convolutional patterns on the endocasts, and identified the corresponding cerebral sulci; for example, in La
Quina, La Chapelle aux Saints, “Sinanthropus”, and “Pithecanthropus erectus”.
In terms of the validity of those inferences, Symington (1916) criticized the simple assumption of thecorrespondence of endocranial morphology to cerebral convolutions and stated that “the simplicity or complexityof the cerebral fissures and convolutions cannot be determined with any degree of accuracy from endocranial
casts.” Le Gros Clarke (1936) generally supported Symington’s view in his paper on the comparison of endocasts
and brains of chimpanzees, and Ogawa et al. (1970) also stated that Symington’s explanations were more plausible
in their study on the Amud endocast. Smith-Agreda (1955) examined more than 300 Modern Human skulls
and found that impressions corresponding to cerebral gyri were observed typically in the anterior and middle
cranial fossa, and to a lesser extent on the inner surface of the lateral wall of the skull. These findings indicate thatthe inference of the cerebral gyri and sulci using the skull might be reliable in the basal portion of the skull but
increasingly difficult towards the vertex.
We have been approaching this issue based on the skulls and brains of extant primate species including human.
First, we confirmed that most of the major sulci were identified on endocasts in Macaca fascicularis (Kobayashi
et al, 2014a). We determined not only the locations of major sulci on the endocasts, but also observed individual
differences of some sulcal patterns. We next analyzed the dry skull specimens of 10 primate species, andcompared the impressions on their endocasts with the sulcal patterns of brains that were so far reported in theliterature. As in the Macaca fascicularis, major sulci were identified on the endocasts in a lemur, most monkeys, anda gibbon although no marked correspondence was observed in a chimpanzee. Moreover, convolutional patterns
were obscured in the parietal region particularly in Macaca fuscata, which have the largest skulls among themonkeys we analyzed. We also observed that endocasts of human infants exhibit more obvious impressions thanin adults although their correspondence to cerebral sulci and gyri is not always clear. These findings indicated that
impressions on the endocasts are more marked in smaller skulls, and are obscured particularly on the dorsolateralsurface in larger skulls, such as adults of greater apes and humans. It prompted us to analyze immature individuals

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of greater apes and humans. We will present the outline of the data that were obtained in collaboration with
Christoph P. E. Zollikofer and Marcia Ponce de León at the Anthropological Institute, University of Zürich.
Concerning the location of the precentral sulcus, Flatau and Jacobsohn (1899) already illustrated the close
relationship between the precentral sulcus and the coronal suture in the macaque, chimpanzee, and human. Evenin lemurs, which lack a homologue of the precentral sulcus, the cytoarchitectonic border obtained in later studies(Brodmann 1909) lies close to the coronal suture. We re-evaluated the relationship between the sulcus and thesuture using modern techniques.
By comparing the CT images of the skull and the brain in Macaca fascicularis, we determined that the lower
limb of the arcuate sulcus and the lower half of the coronal suture were closely located (Kobayashi et al. 2014b).
In other primate species, we confirmed similar relationships but also found that there were some individual andspecies differences. In the modern Japanese, we analyzed the relationship using two different methods. First, wecreated thin-layer silicone endocasts of the calvarias and compared the location of the coronal suture with that ofthe precentral sulcus determined on the brain. The lower portion of the precentral sulcus is located posterior to
the lower half of the coronal suture. It indicates that the caudal border of the prefrontal cortex shifted caudally inthe human compared to the monkeys. To evaluate the accuracy of the silicone endocast method, we also analyzed
2D- and 3D-CT and MR images of Modern Human skulls and brains that were found to be normal on CT and
MR imaging. The results showed relationships between the precentral sulcus and the coronal suture that wereconsistent with the silicone endocast findings. Further evaluation of this approach will be necessary, for example
in greater apes, before we apply it to fossil hominids.
Our findings thus showed both the possibilities and limitations of the inference of the cortical subdivisions basedon the skull morphology. Data on smaller and immature individuals might play a key role in the inference of thebrain structure and functions of fossil hominids.
References
Brodmann K (1909) Vergleichende Lokalizationslehre der Grosshirnrinde in ihren Prinzipien dargestellt auf Grund des Zellenbaues.
Barth, Leipzig.
Flatau E, Jacobsohn L (1899) Handbuch der Anatomie und vergleichenden Anatomie des Centralnervensystem der Säugetiere. 1.
Makroskopischer Teil. Karger, Berlin.
Kobayashi Y, Matsui T, Haizuka Y, Ogihara N, Hirai N, Matsumura G (2014a) Cerebral sulci and gyri observed on macaque endocasts. In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 2:
Cognitive and physical perspectives. Springer, New York, pp. 131-137.
Kobayashi Y, Matsui T, Haizuka Y, Ogihara N, Hirai N, Matsumura G (2014b) The coronal suture as an indicator of the caudal border of the macaque monkey prefrontal cortex. In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in
Neanderthals and Modern Humans, vol. 2: Cognitive and physical perspectives. Springer, New York, pp. 139-143.
Le Gros Clark WE, Cooper DM, Zuckerman S (1936) The endocranial cast of the chimpanzee. Journal of Royal Anthropological
Institute 66: 249-268.
Ogawa T, Kamiya T, Sakai S, Hosokawa H (1970) Some observation on the endocranial cast of the Amud Man. In: Suzuki H, Takai
F (eds) The Amud Man and his cave site. University of Tokyo Press, Tokyo.
Smith-Agreda V (1955). Über die Verteilung der Impressiones gyrorum an der Innenseite des Gehirnschädels des Menschen. Mit
Benützung von Endokranialausgüssen. Deutsche Zeitschrift für Nervenheilkunde 173: 37-68.
Symington J (1916) Endocranial casts and brain form: A criticism of some recent speculations. Journal of Anatomy and Physiology
50: 111-130.

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Inferring fossil brains from endocasts: New methods andresults
Marcia S. Ponce de León1 and Christoph P. E. Zollikofer1*

1 Anthropological Institute and Museum, Unversity of Zurich, Zurich, Switzerland. *Email: zolli@aim.uzh.ch

Human evolution is characterized by two key processes: transition from quadrupedal to bipedal locomotion,and transition from large-faced/small-brained species to small-faced/large-brained species. While evolutionary changes in brain size can be inferred from endocranial volume measurements, evolutionary changes in brain
shape and brain organization are more difficult to track. The principal challenge is to infer the external morphology of the brain from the morphology of the endocranial cavity, and to infer structural and possibly functionalchanges in the brain from its external morphology. Here we present new concepts and methods of endocast-brain
inference. One approach is to use methods of forensic soft tissue reconstruction. While these methods have been
designed to reconstruct “external” soft tissues such as the face, the underlying concepts can also be applied to reconstruct “internal” soft tissues such as the brain. Another approach is to use clinical CT and MRI data to characterize the relationship between the endocranial cavity and the brain in living subjects. Together, these methods
provide new insights into possible structural reorganization of the brain during the course of human evolution.

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Estimating the cerebral and cerebellar volumes of Neanderthals
and Middle and Upper Paleolithic Homo sapiens
Daisuke Kubo1*, Hiroki C. Tanabe2, Osamu Kondo3, Hideki Amano4,
Akira Yogi5, Sadayuki Murayama5, Hajime Ishida6 and Naomichi Ogihara4
1. Faculty of Health and Sport Sciences, University of Tsukuba, Japan; 2. Department of Social and Human Environment, Nagoya
University, Japan; 3. Department of Biological Sciences, The University of Tokyo, Japan; 4. Department of Mechanical Engineering,
Keio University, Japan; 5. Department of Radiology, University of the Ryukyus, Japan; 6. Department of Human Biology and Anatomy,
University of the Ryukyus, Japan. *Email: dddkubo@gmail.com

Introduction
The trajectories of brain size increase in the hominin lineage have been revealed by many fossil studies because
endocranial volume is the good estimator for brain size. Yet, we have only limited knowledge about the fossil
brain substructures due to difficulties to estimate them, including the cerebellar size in the hominin lineage.
Weaver (2001, 2005) reported that the cerebella of Neanderthals and Cro-Magnon 1 were smaller than thoseof the recent Modern Humans. Weaver (2010) discussed the association among the brain, gene and cultural
evolutions in the Late Pleistocene, postulating that the relative cerebellar size did not reach the recent modern
condition until some time after 28,000 years BP.
On the other hand, Kubo et al. (2014) pointed out that the Weaver’s estimation method for cerebellar volume is
not useful to apply to Modern Humans (and probably the close hominin relatives like Neanderthals) due to thenature of the reference sample and likely affected by the inappropriate calibration. Thus, the cerebellar volumes of
Neanderthals and Cro-Magnon 1 have to be re-examined. In addition, because there is no report about cerebellar
volume estimates for earlier Modern Humans older than Cro-Magnon 1, it remains unclear whether the Late
Pleistocene Homo including Neanderthals and early Modern Humans shared relatively small cerebella.
We previously reported a new method to estimate cerebellar volumes from endocasts, which was devised byusing MRI data of a living Modern Human sample (Kubo et al. 2014). This paper reports a method to estimate
cerebral volumes from endocasts, and then calculates the cerebral/cerebellar volumes of fossil specimens including
Neanderthals, Cro-Magnon 1, and Middle Paleolithic Modern Humans. We compare the cerebellar volume
fraction within the whole brain volume among these Late Pleistocene Homo specimens and living humansample in order to assess whether the relatively small cerebellum is an ancestral condition (to the recent Modern
Humans) shared among early Modern Humans and Neanderthals.
Materials and methods
First, we investigated the volumetric relationships between endocranial subdivisions and brain parts in a reference
sample consisting of thirty-two healthy Japanese subjects (18–22 years-old; 16 males and 16 females) in order todevise a method to estimate the cerebral and cerebellar volumes from skulls.

116

We measured the volumes of cerebrum (VCBR), cerebellum (VCBL), endocranial cavity (VEC), and two endocranial
parts (VPCF and VUEC) of the reference sample based on the magnetic resonance imaging (MRI) data withreconstructed voxel resolution of ~1×~1×1 mm. VPCF is synonymous with PCFV in Kubo et al. (2014), which is a
substantial volume of the posterior cranial fossa (PCF) region except for a small portion anterior to the openingof the internal acoustic meatus and basion. VUEC is the volume of the upper endocranial cavity, which excludes
the whole PCF region (i.e. VPCF and the aforementioned small portion). We then calculated the correlation

Looking into the Neanderthal Braincoefficients and reduced major axis (RMA) equations between VCBL and VPCF and between VCBR and VUEC,
respectively.
The RMA equations are primarily to be applied to the volumes derived from computed tomographic (CT) data,
while the VPCF and VUEC values of the reference sample are derived from MRI data, which might have somesystematic error relative to the CT-based measurements. Hence, before calculating the RMA equations, wecalibrated the VPCF and VUEC values of the reference sample by multiplying by factors of 1.002 (for VPCF) and1.014 (for VUEC) respectively, which were the ratios derived from paired comparison between the CT and MRI
measurements of the same individuals (n=3).
We then estimated the VCBR and VCBL of fossil crania by using the RMA equations thus determined. The fossil
sample includes two Middle Paleolithic H. sapiens from Levant (Skhul 5 and Qafzeh 9), an Upper Paleolithic H.
sapiens (Cro-Magnon 1), and four Neanderthals (La Chapelle-aux-Saints, Gibraltar 1, Amud 1, Teshik-Tash).
The VEC, VUEC, and VPCF of Teshik-Tash were taken from the CT imagery of the plaster endocast. Those of theother fossil specimens were taken from the endocranial models which were reconstructed from the CT data of
the original crania. The substantial missing parts of Amud 1 and Gibraltal 1 endocasts were interpolated by thepreserved parts of Gibraltal 1 and La Chapelle-aux-Saints (see Ogihara et al.’s paper in this conference for moredetails).
As a measure of the relative cerebellar size, we also calculated ‘volume fraction of cerebellum’ (FCBL), which isthe proportion of the VCBL within the total brain volume (VTOT). The VTOT estimates are obtained by multiplying
the sum of VCBR and VCBL by a factor of 1.0143, which is derived from the volumetric data of the brain parts(e.g. the medulla oblongata and mesencephalon) of a human cadaver (Stephan et al., 1981). We compared the
VCBL, VCBR, VTOT, and FCBL estimates among four groups (i.e. Neanderthals, Middle Paleolithic H. sapiens, Upper
Paleolithic H. sapiens, and recent Modern Humans) by performing Mann-Whitney U test with Holm correction.
In addition, we calculated the FCBL estimates based on the data of Weaver (2001) for comparison. Weaver (2001)
reported the brain weight estimates of the fossil sample. Hence, we converted the weight values into VTOT by afactor of 1.036 from Stephan et al. (1981).
Results and discussion
The VPCF and VCBL, and the VUEC and VCBR in the living human sample were highly correlated respectively (Pearson’s
correlation coefficient: 0.884 and 0.944). Hence, we can reasonably assume that the two endocranial metrics areuseful estimators for the cerebral/cerebellar volumes as far as the estimation is applied for the fossil specimen
whose endocranial size or shape is not so far from that of the living Modern Humans. Based on the volumetric
relationships, we determined two RMA equations for VCBL and VCBR estimation (Fig. 1, VCBL = 0.775 VPCF +39.5;
VCBR = 0.834 VUEC −27.6).
The FCBL estimates of the four Neanderthal specimens calculated using these RMA equations were significantly
smaller than those of living human sample (Fig. 2, Mann-Whitney U test with Holm correction: p = 0.0055),
while the VCBL, VCBR, or VTOT estimates were not significantly. The FCBL estimate of the Cro-Magnon 1 is small,narrowly within the range of the recent Modern Human variation. The VCBL, VCBR, VTOT, and FCBL estimates of
Skhul 5 and Qafzeh 9 crania were comparable to those of the living human sample. These results suggest that the
Neanderthals had relatively (but not absolutely) smaller cerebella compared to the recent Modern Humans, while
the earlier Middle Paleolithic H. sapiens represented by Skhul/Qafzeh shared relatively large cerebella with therecent Modern Humans.
The results of this study were partly compatible with Weaver (2001, 2005), which reported that the cerebella of
Neanderthals and Cro-Magnon 1 were smaller than the means of the recent Modern Humans. However, our

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FCBL estimates were not as extremely small as the FCBL estimates derived from the data of Weaver (2001) (Fig. 2).
The latter FCBL estimates for the Neanderthal specimens (less than 0.8 on average) are considerably small among
mammalian species (Clark et al. 2001). As we previously detailed (Kubo et al. 2014), the estimation method
devised by Weaver (2001) has some unignorable problems. Therefore, we think that the cerebellar proportion
within the total brain volume of Neanderthals is not so much small as previously reported.
References
Clark DA, Partha PM, Wang SSH (2001) Scalable architecture in mammalian brains. Nature 411: 189–193.
Kubo D, Tanabe HC, Kondo O, Ogihara N, Yogi A, Murayama S, Ishida H (2014) Cerebellar size estimation from endocranial measurements: An evaluation based on MRI data. In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in
Neanderthals and Modern Humans, vol. 2: Cognitive and physical perspectives. Springer, New York, pp. 209–215.
Stephan H, Frahm H, Baron G (1981) New and revised data on volumes of brain structures in insectivores and primates. Folia Primatologica 35: 1–29.
Weaver AGH (2001) The cerebellum and cognitive evolution in Pliocene and Pleistocene hominids. Dissertation, University of New
Mexico.
Weaver AH (2005) Reciprocal evolution of the cerebellum and neocortex in fossil humans. Proceedings of the National Academy of
Sciences, USA 102: 3576–3580.
Weaver AH (2010) Cerebellum and brain evolution in Holocene humans. In: Broadfield D, Yuan M, Schick K, Toth N (eds) The human brain evolving: Paleoneurological studies in honor of Ralph L. Holloway. Stone Age Institute Press, Gosport, IN, pp. 97-105.

Fig. 1 Biplots of the volumes of the endocranial and brain parts obtained from
the MRI data of the living human subjects. Left: the posterior cranial fossa (VPCF)
and cerebellum (VCBL); Right: the upper endocranial parts (VUEC) and cerebrum(VCBR). Red solid lines represent the reduced major axis (RMA). Black solid lines
represent the ordinary least squares (OLS) regression, and dotted and dashedlines are the 95 % confidence and prediction intervals of OLS regression.

Fig. 2 Estimates of ‘volume fraction of cerebellum’ (FCBL), the proportion of
the cerebellar volume within the total brain volume. The results of this study
are shown in the left four columns. The comparative estimates calculatedusing the data from Weaver (2001) are shown in the right end column. The
mean FCBL of the recent Modern Human sample of this study is shown as
a horizontal line, with comparison to that of Weaver (2001) represented by
the open circle in the right end column.

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Endocasts of Neanderthals and early Modern Humans:
Virtual reconstruction and geometric morphometric analyses
Naomichi Ogihara1*, Hideki Amano1, Yusuke Morita1, Osamu Kondo2,
Daisuke Kubo3, Hiromasa Suzuki2, Takashi Michikawa4, Masaki Moriguchi5,
Hajime Ishida6 and Masato Nakatsukasa7
1. Department of Mechanical Engineering, Keio University, Japan; 2. University of Tokyo, Japan; 3. University of Tsukuba, Japan; 4.
Osaka University, Japan; 5. Chuo University, Japan; 6. University of the Ryukyus, Japan; 7. Kyoto University, Japan.
*Email: ogihara@mech.keio.ac.jp

Introduction
In order to examine the possible differences in learning ability between Neanderthals and early Modern Humans
in terms of the brain anatomy, original antemortem appearance of fossil crania that enclose the brains must be
correctly restored. In the present study, we performed the virtual reconstructions of three Neanderthal crania,
Amud 1, Gibraltar 1, and Chapelle-aux-Saints 1, and three early Modern Human crania, Qafzeh 9, Mladec 1,
and Cro-Magnon1. The endocranial morphology was then analyzed using geometric morphometrics.
Materials and methods
In the case of Amud 1, we mathematically reassembled the fragments based on smoothness of the joints amongfragments (Kikuchi and Ogihara 2013). As a result, smooth yet globally consistent assembly of the fragments ofAmud 1 cranium became possible. However, the cranial base and endocranial surface of Amud 1 are largely missing or damaged. We therefore warped the Gibraltar 1 and La Chapelle-Aux-Saints 1 onto the Amud 1 crania toestimate the missing basicranial and damaged endocranial regions. In the case of Gibraltar 1, the La ChapelleAux-Saints 1 was warped to compensate for the missing regions, and vice versa in the case of La Chapelle-AuxSaints 1. Reference Modern Human cranium was then warped onto the new reconstruction by iterative thinplate spline deformation to compensate for the missing parts and complete exo- and endocranial morphology
was generated. The endocasts of the early Modern Humans were also generated in the same manner.
We then analyzed the morphological variability of the endocranial shape of the fossil skulls using landmarkbased geometric morphometrics. On the endocranial surface, a total of 16 anatomical landmarks were digitized.
Equally-spaced points along the anterior boundary of the anterior cranial fossa and the inferior border of thegroove for transverse sinus were also defined as landmarks (Morita et al. 2013). We define sliding semi-landmarkson the entire endocranial surface based on the shortest paths connecting pairs of anatomical landmarks andequally-spaced points along the midsagittal curve. Therefore, the variability in endocranial shape was examinedbased on a total of 171 anatomical and sliding semi-landmarks. We also included endocasts of the modern Japanese and European populations housed at Kyoto University and the University of Tokyo, respectively, for comparisons.
Results and discussion
Our results demonstrated that exo- and endocranial shapes are quantitatively different between Neanderthals
and Modern Humans. Specifically, our geometric morphometric analyses revealed that Modern Human cranium
shows relative enlargement of the cerebellar region, and relative expansion of the parietal area as suggested byother studies (Bruner et al. 2010; Gunz et al. 2010; Weaver 2005), possibly indicating that neuroanatomical organization may be different between the two species. Recent morphological studies on the pattern of endocranial

119

Looking into the Neanderthal Brainontogeny of Neanderthals and Modern Humans suggested that relative expansion of the parietal and cerebellar
regions are observed during the early postnatal period in Modern Humans so that the endocranium becomes
more globular, but it is not the case in Neanderthal lineage (Gunz et al. 2012). Ontogenetic differences of the
cranium and hence the brain between Neanderthals and early Modern Humans may thus differentiate developmental process of learning and social skills in early childhood, possibly leading to a critical difference in innate
learning capacity between the two populations.
References
Bruner E, Martin-Loeches M, Colom R (2010) Human midsagittal brain shape variation: Patterns, allometry and integration. Journal of Anatomy 2010: 589-599.
Gunz P, Neubauer S, Golovanova L, Doronichev V, Maureille B, Hublin JJ (2012) A uniquely Modern Human pattern of endocranial development. Insights from a new cranial reconstruction of the Neandertal newborn from Mezmaiskaya. Journal of Human Evolution 62: 300-313.
Gunz P, Neubauer S, Maureille B, Hublin JJ (2010) Brain development after birth differs between Neandertals and Modern Humans.
Current Biology 20: R921-R922.
Kikuchi T, Ogihara N (2013) Computerized assembly of neurocranial fragments based on surface extrapolation. Anthropological Science 121: 115-122.
Morita Y, Ogihara N, Kanai T, Suzuki H (2013) Quantification of neurocranial shape variation using the shortest paths connecting
pairs of anatomical landmarks. American Journal of Physical Anthropology 151: 658-666.
Weaver AH (2005) Reciprocal evolution of the cerebellum and neocortex in fossil humans. Proceedings of the National Academy of
Sciences, USA 102: 3576-3580.

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Exploring the difference of brain anatomy and function between
Neanderthals and Modern Humans: Neuroanatomical andfunctional neuroimaging approach
Hiroki C. Tanabe1*, Takanori Kochiyama2, Norihiro Sadato3 andNaomichi Ogihara4

1. Graduate School of Environmental Studies, Nagoya University, Japan; 2. Department of Cognitive Neuroscience, Brain Information
Communication Research Laboratory Group, Advanced Telecommunications Research Institute International, Japan; 3. Department of
Cerebral Research, National Institute for Physiological Sciences, Japan; 4. Faculty of Engineering, Keio University, Japan.
*Email: htanabe@lit.nagoya-u.ac.jp

Introduction
Conventionally, the differences in learning ability between H. sapiens and H. neanderthalensis have been
evaluated in terms of skull shape. Such evaluations originally aimed at exploring the differences in activity ofthe brain contained in the skull. However, previous studies either created endocranial casts from reconstructedskulls to compare morphological features of the two species, or extrapolated differences in brain shape fromskull landmarks for the discussion of functional distinctions. Most of the results were approximate volume orregional shape calculations based on cranial or skull shape, and no research efforts have been made to detect
precise functional differences by superimposing cranial images of one species on those of the other within one
framework. In addition, previous studies have not fully established the relationship between cranial morphology
and the anatomical localization of brain function.
As magnetic resonance imaging (MRI) techniques have accelerated functional and anatomical brain imaging andthe non-invasive direct measurement of living human brain activity (i.e. functional MRI) (Huttel et al. 2009), weare able to explore the functional localization of living humans (Friston 1997). During the last 30 years, methods
used for the analysis of these data have also been developed. Here, we apply the anatomical transformation andstandardization method used in functional brain imaging research to the morphological estimation of the cerebral
parenchyma from reconstructed Neanderthal’s skull. This approach will allow for high-precision reconstructionof the fossil brain, as well as a direct comparison of brain morphology between ancient and modern humans.
Moreover, we refer the functional brain map in the Modern Humans, corresponding to the anatomicallydifference regions.
Framework of the analysis
The method of computational neuroanatomy is able to perform quantitative analysis of the biological shape ofthe brain (Ashburner and Friston 2007). This methodology has been developed in parallel with the analysis offunctional brain mapping that measures living human brain activity (Huttel et al. 2009). The image registration
technique uses spatial transformation of the images. Among the various spatial transformation techniques,a promising one that has recently been developed is the large deformation diffeomorphic metric mapping(LDDMM) approach. The LDDMM approach of high-dimensional non-linear transformation has several
million degrees of freedom, as opposed to the one thousand in conventional non-linear transformation. Here,we employed DARTEL algorithm (Ashburner 2007), which is one of the LDDMM approach and is popularin the functional neuroimaging research field. As standardized stereotaxic space such as Montréal Neurological
Institute (MNI) space is available using DARTEL (i.e. the image can be transformed into this space by adding
a few steps), we can comparably easily integrate the estimation and creation of a Neanderthal ‘fossil brain’ withfunctional neuroimaging studies of Modern Humans.

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Looking into the Neanderthal Brain

The strategy is as follows: First, the skull shape of reconstructed computed tomography (CT) images of the
Neanderthal is spatially deformed to the Modern Human skull shape segmented from the MRI image. TheDARTEL algorithm is used to estimate a globally one-to-one smooth and continuous mapping between theseskull shapes. Using the deformation or flow field in DARTEL, the Modern Human cerebral cortex segmented
from the MR image is then inverse-transformed to construct the virtual Neanderthal’s cortex, which isconsequently best fitted to its skull shape. If this strategy goes well, we can use computational morphometry. We
used deformation-based morphometry (DBM) (Ashburner et al. 1998; Chung et al. 2001; Chung et al. 2003)
and automated anatomical labelling parcellation based morphometry. We used Amud 1, Giblarter 1, and La
Chapelle-aux Saints 1CT images as the samples of Neanderthals, whereas more than 500 Modern Human MRimages were used to direct comparison.
The important point of the functional neuroimaging research is that the results of the functional MRI only show
the correlation between a specific task or task condition and the brain activity corresponding to the task, so that
there is no causal relationship. We could not conclude the exact relationship between specific brain regions andmental processes or functions. Therefore, we have to perform similar functional MRI experiments and collectevidence about which part of the brain is truly correlated to the specific function. Meta-analysis is a tool for
realizing this. The goal of the meta-analysis is to show that the consistently activated regions in a set of studies
are related to the same psychological or mental process in a quantitative or statistical manner (Warger et al.
2007). Fortunately, almost all neuroimaging studies are spatially transformed to a standardized stereotaxic space.
If we can select suitable functions of the human brain, the map corresponding to the differences in the cognitive
abilities between H. neanderthalensis and H. sapiens can be elucidated. Here, we performed meta-analysis ofseveral cognitive functions those might be related the difference between them.
Results and discussion
The results of DBM and AAL based morphometry showed that the parietal regions and hemispheres of
cerebellum were larger in Modern Humans rather than in Neanderthals, whereas widespread occipital regions
were larger in Neanderthals rather than in Modern Humans. Recently, many studies demonstrated that
hemispheres of cerebellum play an important role for not only motor-related function but also higher cognition
such as language, working memory, social abilities. Meta-analysis also showed that the lateral cerebellum is
activated during social cognitions such as mentalizing, perspective taking tasks. Unlike anatomical complex
neuronal network in the cerebrum, neural circuit of the cerebellum is rather simple. A functional unit contains
500 Purkinje cells and there are approximately 5,000 units in the Modern Human cerebellum. Recent functional
neuroimaging studies demonstrated that the cerebellum has a variety of inner models that reproduce the dynamic
essential properties of mental representations, which is deeply related to ‘learning’ (Ito 2008). As larger cerebellum
is able to contain many inner models, Modern Humans can have more mental representations. In addition, lateralhemisphere of the cerebellum has rich connection to frontal and parietal regions of the cerebrum. The present
results indicate that Modern Humans use brain more efficiently using larger cerebellum, which maybe one of thecause of replacement.

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References
Ashburner (2007) A first diffeomorphic image registration algorithm. Neuroimage 38: 95-113.
Ashburner J, Friston KJ (2000) Voxel-based morphometry-the methods. Neuroimage 11: 805-821.
Ashburner J, Hutton C, Frackowiak R, Johnsrude I, Price C, Friston K (1998) Identifying global anatomical differences: Deformation-based morphometry. Human Brain Mapping 6: 348-357.
Chung MK, Worsley KJ, Paus T, Cherif C, Collins DL, Giedd JN, Rapoport JL, Evans AC (2001) A unified statistical approach to
deformation-based morphometry. Neuroimage 14: 595-606.
Chung MK, Worsley KJ, Robbins S, Paus T, Taylor J, Giedd JN, Rapoport JL, Evans AC (2003) Deformation-based surface morphometry applied to gray matter deformation. Neuroimage 18: 198-213.
Friston KJ (1997) Analyzing brain images: Principles and overview. In Frackowiak RSJ, Friston KJ, Frith CD, Dolan RJ, Mazziotta

Looking into the Neanderthal Brain

JC (eds) Human brain function, 1st edn. Academic Press, Massatusetts, USA.
Huttel S, Song AW, McCarthy G (2009) Functional magnetic resonance imaging, 2nd edn. Sinauer Associates Inc. Massachusetts,
USA.
Ito M (2008) Control of mental activities by internal models in the cerebellum. Nature Reviews Neuroscience 9: 304-313.
Warger TD, Lindquist M, Kaplan L (2007) Meta-analysis of functional neuroimaging data: Current and future directions. Social
Cognitive and Affective Neuroscience 2: 150-158.

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The RNMH project: A summary

Yoshihiro Nishiaki

The University Museum, The University of Tokyo, Japan. Email: nishiaki@um.u-tokyo.ac.jp

The RNMH project posed an explicit research goal to investigate the replacement of Neanderthals by Modern
Humans. This is to test the “learning hypothesis,” a working hypothesis surmising that the replacement occurred
as a result of differences in learning (abilities/strategies/behaviors) between the two populations. The project
was launched in 2010 to run for five years. It involves six main groups of researchers representing prehistoricarchaeology, cultural anthropology, theoretical anthropology, environmental science, physical anthropology, andneuroscience. The researchers participating in this project number almost 70, including those specialized in otherrelated disciplines. While the research is expected to continue into the next stage, the targets and achievements ofthe project thus far are summarized here for evaluation on the occasion of this international meeting.

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Closing Remarks

Neanderthals and Modern Humans: The contributions of theRNMH project
Ofer Bar-Yosef

Department of Anthropology, Harvard University, USA. Email: ofer.baryosef@gmail.com

Originally Neanderthals were a European population that emerged some 400-200,000 years ago from an earlier
population. They were successful in surviving through several glacial periods whether across temperate or Mediterranean Europe. They were also effective travellers who migrated eastwards reaching western and central Asia.
They formed a meta-population archaeologically represented by fossil remains and numerous sites and assemblages. Few fossils produced the aDNA information indicating a degree of homogeneity among the people. The
archaeological remains of their past life ways exposed a few burials and isolated bones and teeth in many sites, aswell as food refuse (including animal bones and as yet rare plant remains, often due to poor preservation). Wellpreserved anthropogenic deposits, such as in the Mediterranean basin, hearths were uncovered as well as middens
suggesting acts of cleaning the living surfaces. Climatic fluctuations had only minimal effects and excavated sties
in northern latitudes indicate their special physical adaptations that apparently contributed particular genes toModern Humans with whom they interbred in various locations.
We are able to trace the presence, movements and choices of sites of Neanderthal groups through the analysis of
the abundant lithic assemblages. Such geographic tracing is done through the detailed reconstructions of various operational sequences that begins from the search and choice of raw materials, the preliminary preparationof the nodules, the shaping of the cores, detachment of blanks, choice of the selected few for use and/or further
retouching, and discard. The rejected cores that were ‘exhausted’ in the eyes of the local knappers and thus considered useless, could have serve as toolstones the children of the group. We note that the assumption based on
the experience of flint knappers is that teaching the skills of making stone tools begins at an early age. The process could have been similar to teaching children when they grow up how to use chopsticks as the parents speak
and demonstrate the gestures in their own language. Similarly kids could learn how to make stone tools throughobservations and imitation, following instructions and encouragements by the adults. In brief, the education at
an early age is the key for figuring the use of each operational sequence. This was the main target of the RNMH
project, directed by Prof. T. Akazawa and his associates.
At the time when the western world was discussing and debating issues such as the chronology of Neanderthals’sites, the markers of their cultures, their diets (either mostly based on meat or just mixed), issues of longevity,
health, behavior under stress conditions and their cognitive development, the RNMH project took essentailly adifferent direction. Issues of the evolution of the brain, differences between human populations, social relationships and basic were not abandoned. But in my view, the main contribution of this project was by raising theplace of learning, teaching and transmission among past societies to the top of the list of our concerns in understanding the behavior of humans during the Paleolithic. Thus, not only Neanderthals were examined but also
Modern Humans. And because the main archaeological finds are stone tools and animal bones, to which we can
now add the study of microscopic residues, I personally felt that the RNMH project is heading in the right direction. However, as I was originally trained by the late F. Bordes and was already biased thinking that different
groups within the same meta-population would have their own set of operational sequences for the productionof stone tools employed for various tasks. Microwear and edge damage demonstrate that various tool types were
employed for the same activities (e.g., butchering, whittling, wood working, etc.). Thus the RNMH project attracted my attention.

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Closing Remarks

Here I would like to mention that interpreting the known Paleolithic knapping techniques, assemblage composition and tool types are today not exclusively the domains of archaeologists. The reports produced by archaeologists who painstakingly classify all the uncovered elements collected during the course of surveys and excavations,became sources for ideas and interpretations by psychologists, brain scientists, researchers of cognitive evolution
and more. Lively discussions concerning the interpretation of “modern behavior” of particular prehistoric culturalcontexts continued although it became clear in recent years that humans at different times in various regions
produced particular and sometimes unique, material culture elements. In some cases “modern behavior” was identified by prevalence of blade production, body decorations (beads and pendants made of different raw material),rock art, intra-caves murals and open-air sites engravings, as well as mobile imagery (e.g., figurines, decorated organically made artifacts). But in more than one region many of these cultural elements are missing but the people
themselves as shown by the morphology of their skulls, were identified as ‘modern’. Even the efforts to trace thepresence of language and symbolic behavior as currently incorporated within the enhanced working memory, wasproven to be a tough endeavor.
Indeed, the RNMH project directed a portion of the time and funds into the investigations of social learning,teaching, and creativity among children including hunting and gathering societies. Recording the differences
and similarities among individuals, and limitations to human behaviors imposed by social learning were also researched. As expected, some needed testing was carried out among hunting and gathering societies. The insights
gained from analyzing teaching and learning among several groups of children of foragers concerning their activities are used in order to consider their resilience, and how they face demanding situations. These studies corroborate what is achieved through the investigations of other Modern Humans when they asked to face variablesocial situations or in testing their foresights. The cases studied and reported today are based on Modern Humanbehavior but several evolutionary hypotheses take into account the contributions from primate behavior, but unfortunately we miss the behaviors of Homo erectus populations. Thus the challenge to find if there were fundamental differences among ‘cultural learning’s’ between early Modern Humans and Neanderthals’ longer history is adaunting task. The debate often centers on ‘problem-solving’ and ‘creativity’ between these two meta-populations.
Various analyses demonstrate a greater degree of conservatism among the Neanderthals possibly due to the fragmentation of their Eurasian populations. In spite of being successful foragers and migrants they were taken overby Modern Humans. It seems that the an increased degree of social flexibility, improved foresights, better meansof communication, and perhaps having a more efficient technology, facilitated Modern Humans to take over thisthe Eurasian continent within no more than 10,000 years and perhaps much less.

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Closing Remarks

Poster Presentations

Past and present research on the Middle and Upper
Paleolithic of Uzbekistan
Otabek Aripdjanov

State Museum of History of Uzbekistan, Uzbekistan. Email: otabek_ar@hotmail.com

Researchers’ great interest in studying the Paleolithic of Uzbekistan is based on their belief that this region
represents a contact zone, or rather some kind of a “transit place” for ancient human populations occupying Eastand West Asia and other parts of the Paleolithic world. To date, Paleolithic research of the region has yielded asufficient amount of archaeological materials. Analysis of this record allows us to track the evolution of the mostprimitive populations and their cultures and raise questions on the origins of Paleolithic communities living in
the territory of what is now modern Uzbekistan.
The first discoveries of Paleolithic cultural remains in Uzbekistan were made in the 1930s during fieldwork bygeologists and archaeologists at a number of locations. Between 1931 and 1934, G. V. Parfyonov found a series ofcaves and rockshelters; he partially excavated Machay Cave in the Shirabaddarya Valley (Surkhan-Darya region).
This site was investigated by U. Islamov again in the 1970s, when he revealed stone and bone tools as well as
human remains of the Mesolithic period.

P1

The archaeological records from the Uchtutin Bukhara region comprise the most complete chronologicalsequence starting from the Lower Paleolithic period. Long-term research there has yielded abundant materials
from the Acheulian, Mousterian, Upper Paleolithic, and Neolithic periods. In the Akhangaron River valley of the
Tashkent region is the unique multi-level Paleolithic site of Kulbulak, the focus of long-term research since 1963.
Rough stone tools such as cores and massive flakes from slate and flint were found there. In addition, Kulbulak
remains the only known Mousterian site in the region.
The main level of the multi-level cave of Sel-Ungur (Fig. 1) in the Fergana Valley (700–500 ky BP) also belongs
to the Lower Paleolithic. This site was for many years studied by the academician U. Islamov. Excavations there
yielded disc-shaped cores, unifaces and bifaces, broad flakes, scrapers, knives, and choppers made on pebbles.
Moreover, sensational finds included teeth, a humerus, and the back part of a skull of the anthropoid called“Fergantrop” (Fig. 6).
A group of sites from the Mousterian period were found in the Surkhan-Darya region, Chirchik-Akhangara
Valley, Bostanlyk area (Tashkent region), and the Fergana Valley. These groups differ regarding the nature ofprocessing stone and the composition of stone processing tools. One of the most significant sites is Teshiktash
Cave (Fig. 2), discovered by L. P. Okladnikov in 1938. He found not only Mousterian stone tools, but thefirst remains of a Neanderthal child burial ever recorded in Central Asia. Based on the materials recovered, anumber of features of this primitive people’s life were established. Excavation of the site revealed the bones ofsuch animals as goat, deer, and bear. Stone tools were made on-site from quartz, quartzite, jasper, or siliceous
limestone. Tool production technology is typical for the Mousterian, as seen in a disc-shaped core. The main tools
are scrapers and small chisels. The most important discovery was the burial containing the remains of a boy; thisNeanderthal individual’s shape was reconstructe by the famous anthropologist and archaeologist named M. M.
Gerasimov. The grave contained mountain goat horns, reflecting the origin of Neanderthal cult representations.

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In 1939, an Upper Paleolithic site in the city park area of Samarkand was opened by N. G. Kharlamov. It was

Poster Presentations

later investigated by M. V. Voyevodsky. Excavations were renewed in 1958 by D. N. Lev, and then continued by M.
D. Dzhurakulov. In 1960 and 1964, H. K. Nasretdinov found and excavated the Late Paleolithic Ak-Tash Cave.
Numerous stone artifacts of the Mousterian, Upper Paleolithic, and Mesolithic were obtained from the site ofBozsu 1-2. The Mousterian localities of Bozsu 3-5 and Zakh were subsequently discovered.
One of the first Mousterian finds in the Tashkent region was in 1942 by R. V. Smirnov. In 1958, P. T. Konoplya
discovered the Mousterian site Kuprikbosh. The cave of Hodjakent-1 was excavated in 1958–1959 under the
direction of A. P. Okladnikova, and Hodjakent 2 was studied by H. Nasretdinov.
R. K. Suleymanov studied one of the richest cave sites, Obi Rakhmat (Fig. 3), which holds a special place
among Mousterian sites of the Tashkent region. This is because its cultural layers contain many thousands ofstone artifacts in-situ (Fig. 4). For this region, this site is unique because most Paleolithic sites have disturbed
stratigraphic contexts or contain small lithic collections. Throughout the second half of the 20th and beginningof the 21st centuries, analyses of the Obi Rakhmat collections have revealed several cultural and chronologicalphases of development of the regional Paleolithic. Thus, depending on the excavation methods and analysis oflithic materials by researchers, various conclusions have been drawn regarding the chronological and culturalinterpretations of complexes from this site.
In Layer 16 of Obi Rakhmat, six teeth and about 150 small skull fragments (Fig. 5) belonging to a child aged
9–12 years were found in 2003. The scholars M. Glantz, B. Viola, and T. Chikisheva believe that the small skull
fragments have no diagnostic characteristics, indicating the duplication of bones. Regarding the teeth, their large
size (characteristic of Neanderthals or a robust Homo sapiens) and root development provide no information on
the types of hominin. Skull fragments reflect a morphological structure more similar to a Modern Human, witha big parietal lobe uncharacteristic for Neanderthals. The temporal bone also looks modern but the inner ear
labyrinth has no clear morphology. Therefore, there is a mosaic of Modern and Neanderthal types.
In 2000, the southern slope of the Karatyubinsk massif (main watershed valleys of the Kashkadarya andZarafshan rivers) was examined by the archaeologist R. K. Suleymanov. He was in the Ayakchisaya Valley,
where the Middle Paleolithic site of Angilak Cave is located. Under the guidance of R. K. Suleymanov and M.
Glantz, archaeological work was conducted between 2001 and 2004, during which it was revealed that the stone
industries of Angilak were similar in technology and typology to those of Kuturbulak and Amankutan caves.
However, they differ in terms of their strongly amorphous nature and the small sizes of cores, chips, and tools.
Excavations at this site were resumed in 2013–2014 by the State Museum of History of Uzbekistan and the
University Museum of the University of Tokyo under the guidance of R. K. Suleymanov, O. Aripdjanov, and Y.
Nishiaki.
Recent studies of Middle and Upper Paleolithic complexes in Uzbekistan allowed us to specify characteristics ofPaleolithic complexes from certain sites, and to revise the general provisions concerning the Upper Paleolithicof the region. This region became a zone through which humans moved in later times, based on finds fromnumerous Acheulean and Mousterian sites. All analyzed industries are characterized by a clearly establishedcombination of Middle and Upper Paleolithic attributes.

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Poster Presentations

Fig. 1 Archaeological expedition at
the entrance to Selungur Cave (Lower
Paleolithic), 1985.

Fig. 2 General view of Teshiktash Cave (Middle Paleolithic), Surkhandarya
region.

Fig. 4 Stone artifacts of Obi Rakhmat (Middle-Upper Paleolithic transitional
period). Tashkent region.

Fig. 3 General view of Obi Rakhmat Cave,
Tashkent region.

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Poster Presentations

Fig. 5 Human fossils of the Middle Paleolithic from Obi Rahmat, Tashkent
region.

The Neanderthal occupations at Dederiyeh Cave, Syria

Yoshihiro Nishiaki1*, Yousef Kanjou2 and Takeru Akazawa3

1. The University Museum, The University of Tokyo, Japan; 2. National Museum of Ethnology, Japan; 3. Research Institute, Kochi
University of Technology, Japan. *Email: nishiaki@um.u-tokyo.ac.jp

The Middle Palaeolithic sequence of Dederiyeh Cave, northwest Syria, is unique in its association with a seriesof Neanderthal fossil remains from stratified contexts (Akazawa and Muhesen 2002; Nishiaki et al. 2011). One
particular area (Chimney area) of this large cave attests to the continuous occupations of Neanderthals during
the late Levantine Mousterian (ca. 70–50 ka), providing an opportunity to investigate chronological changes in
Neanderthal behaviors. With reference to lithic assemblages and other evidence from this area, excavated between
1989 and 2002, we address how Neanderthal settlement patterns and population size might have changed.
The available data suggest a significant increase in occupational intensity during this time period (Nishiaki et al.
2012). This is demonstrated by a rapid increase in the number and size of fire hearths and a remarkable increasein lithic artifacts per volume of sediment (m3) through time. Furthermore, technological and typological traits
thought to indicate more sedentary occupations also became more remarkable in the upper layers (see Parry and
Kelly 1987). They include the higher frequency of expedient tools at the expense of standardized tools such as
Levallois points and side scrapers, and the more common practice of expedient core reduction rather than carefulcore preparation.
These changes collectively point to a shift from a more ephemeral to more prolonged use of the cave. In otherwords, the base-camp character of the Dederiyeh Cave became more conspicuous in upper layers. This can beinterpreted as responses to the increasingly favorable environmental conditions later in the Mousterian sequenceat Dederiyeh Cave: wild boar, aurochs, and deer indicative of temperate and humid environments appeared in
faunal assemblages of the upper layers, replacing goat remains that indicate colder and drier environments (Griggo
2002). Evidence in the sedimentological records of decreasing soil inflow from outside the cave has also beeninterpreted to show increased vegetation due to increased precipitation (Oguchi and Fujimoto 2002).

P2

Research in the southern Levant highlighted a comparable trend for the entire Levantine Mousterian sequence,in which occupations intensified from the middle Levantine Mousterian onwards (ca. 130 ka). This trend has
been interpreted to indicate a residential shift from a more mobile to a more sedentary, territorial residential
pattern due to population increase (Hovers and Belfer-Cohen 2013). In this vein, findings from Dederiyeh
might suggest that the trend, i.e., demographic increase, continued throughout the late Levantine Mousterian. If
so, future research can utilize the stratified lithic assemblages from Dederiyeh to test theoretical premises often
predicted on the relationship between demography and cultural evolution among human societies; for example,
whether and how a positive correlation between population increase and cultural complexity also existed among
Neanderthal societies.
References
Akazawa T, Muhesen S (2002) The Neanderthal burials: Excavations of the Dederiyeh Cave, Afrin, Syria. International ResearchCenter for Japanese Studies, Kyoto.
Griggo C (2004) Mousterian fauna from Dederiyeh Cave and comparisons with fauna from Umm el-Tlel and Douara Cave.
Paléorient 30(1): 149–162.
Hovers E, Belfer-Cohen A (2013) On variability and complexity: Lessons from the Levantine Middle Paleolithic record. Current Anthropology 54(S8): S337-S357.

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Nishiaki Y, Kanjo Y, Muhesen S, Akazawa T (2011) Recent progress in Lower and Middle Palaeolithic research at Dederiyeh
Cave, northwest Syria. In: Le Tensorer J-M, Jagher R, Otte O (eds) The Lower and Middle Palaeolithic in the Middle East andneighbouring regions. Etudes et Recherches Archéologiques de l’Université de Liège, ERAUL 126. Université de Liège, Liège,
pp. 67–76.
Nishiaki Y, Kanjo Y, Muhesen S, Akazawa T (2012) Temporal variability of Late Levantine Mousterian assemblages from
Dederiyeh Cave, Syria. Eurasian Prehistory 9(1/2): 3–27.
Oguchi T, Fujimoto K (2002) The sediment and paleoenvironment of the Dederiyeh Cave. In: Akazawa T, Muhesen S (eds)
The Neanderthal burials: Excavations of the Dederiyeh Cave, Afrin, Syria. International Research Center for Japanese Studies,
Kyoto, pp. 33–61.

Fig. 1 Mousterian tools and debitage from the upper layers of the chimney area.

132

Fig. 2 Mousterian tools and debitage from the lower layers of the chimney area.

Poster Presentations

The emergence of the Upper Palaeolithic in the Japanesearchipelago
Hiroto Nakata

Aoyama Gakuin University, Japan. Email: hiroto.nakata@gmail.com

The replacement of Neanderthals by Modern Humans can be explained not only by the differences in theevolutionary history of human populations but also by the differences in the nature of society and culture thatthey have experienced. It is important to investigate the causes of the replacement from such a viewpoint as well.
When Neanderthals were replaced by Modern Humans, various cultural/social changes occurred in Eurasia.
Given these archaeological changes, it is necessary to examine each of the regionally diverse cases. In thisperspective I discuss how Upper Palaeolithic cultures formed in the Japanese archipelago.
When we discuss the emergence of the Upper Palaeolithic in the Japanese archipelago, it is important to examine
three aspects: stratigraphy, dating, and stone tool types. The earliest Upper Palaeolithic assemblages are found in
the central part of Honshu, the mainland of Japan. For example, the assemblages in the Ashitaka-Hakone area,
dated to 38-36ka BP, indicate that the earliest inhabitants settled into the Japanese archipelago by at least 40ka.
These early assemblages are divided into several industries. One is the small flake industry which is characterizedby light retouched flakes, scrapers, and drills. There are few formal tools in this industry. The second is thedenticulate industry characterized by denticulates on flakes, picks, and scrapers with inverse retouch. The third
is the pebble tool industry or heavy duty tool industry characterized by choppers, chopping tools, “hand axes”.
Their chronological sequence or functional relationship is currently unclear due to the scarcity of relevant records.
Importantly, these early industries do not include blade technology or polishing technology, which subsequently
characterize the Upper Palaeolithic period of Japan.

P3

Blade technology appears for the first time in Japan around 36-35ka. In Happusan II site, Nagano Prefecture,
blades are detached from unidirectional narrow faceted cores. The blade blanks are then used for the production oftools including pointed blade, burins, and scrapers. The assemblage has another distinctive feature of the bladelet
production from core edge flakes or flake cores. In this initial phase of the blade/bladelet technology, trapezes
and edge ground axes also emerged. Because trapezes or edge ground axes did not develop in China, Korean
peninsula, or Siberia, these lithic technologies probably emerged independently in the Japanese archipelago.
The trapezes may have derived from preceding small flake industry while it is difficult to trace the origin of theedge ground axes. The form of the edge ground axe is similar to large pointed tools, classified as “hand axes”, in
Kanadori site, and it is technologically similar to chopper/chopping tools using flat pebble stones. The use-wear
analysis shows that the edge-ground axes have been used for cutting trees and modifying wood. Thus, it may havederived from previous large cutting tools. However, it is impossible to trace the origin of polishing technology
used for the production of this lithic type.
During 34ka-32ka, a very distinctive settlement organization, named “Kanjou burokku”, appeared throughout
Honshu, the main island of Japan. This consists of several lithic clusters arranged in a circular pattern, whichappears to have occurred exceptionally in the Japanese archipelago. Although this settlement type used to beconsidered as representing aggregation sites for the big game hunting, it provided at least opportunities forthe exchange of information among the groups that gathered at the site. Such social networks possibly explain
the emergence of standardized tools, such as blades, trapezes, and edge ground axes as well as their regional

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Poster Presentations

diversification.
The following summarizes the above descriptions regarding the emergence of the Upper Palaeolithic in theJapanese archipelago.
(1) Widely accepted archaeological records indicating the colonization of the Japanese archipelago date after
40ka. The earliest industries basically consist of the small flake industry and the heavy duty tool industry. Because
these lithic technologies are similar to the Early Palaeolithic industry in eastern Eurasia, it would be suggested
that the lithic industry before blade technology in Japan can be related to the east Eurasian lithic industry.
(2) Major changes have appeared after 36ka. One is the formation of trapeze industry. This industry was fully
equipped with various types of trapezes. This trapeze industry may have derived from the preceding small flakeand denticulate industries. Edge ground axes also appeared.
(3) Another change is the formation of blade technology. Large blades and bladelets were produced from narrow
faceted cores, and the blade blanks were then modified into pointed blades, scrapers, and burins. There are twodifferent views regarding the origins of the blade technology in Japan. One suggests that the blade technology
was introduced to the Japanese archipelago from adjacent areas, such as Siberia and China, whereas the otherargues that the blade technology developed independently in Japan from previous lithic industries. The former
scenario currently appears to have gained more supportive records.
(4) After 34ka, the blade technology became more standardized. Blades of this phase are manufactured fromexotic raw materials of higher quality than those produced from local materials. The blade technology of thisphase is characterized by prismatic cores and core debitage such as core rejuvenation flakes, striking-platform
rejuvenation flakes, surface rejuvenation flakes, and core edge trimming chips. Bladelets are detached from coreson blades and piece esquillees. In this phase, some stone tool types became more standardized. They are produced
from particular raw material types or blank types.
(5) Archaeological evidence for symbolic behaviors are scarce until 30ka. It has become prevalent since the LGM
period in Japan.
In this paper, the beginning of the Upper Palaeolithic in Japan is defined by the emergence of trapezes and edge
ground axes. Therefore, preceding lithic industries should be separated from the Upper Palaeolithic. In eastern
Eurasia, the Palaeolithic culture is usually divided into two periods, the Early and Late (Upper) Palaeolithic,
instead of the tripartite periodization in western Eurasia. Based on the two-period system, the lithic industries
before blade and trapeze assemblages are grouped together in the Early Palaeolithic. However, these industries
are few and their chronological or functional relationship is unclear. We also need to carefully evaluate theirculture-historical links to the subsequent blade and trapeze industries. In addition, even if the lithic industries
before the trapeze or blade technologies are included in the Early Palaeolithic, it does not mean that their makers
are not Homo sapiens.
In the case of the Japanese archipelago, the transition to the Upper Palaeolithic is regionally diverse. This paper
focused on the records from the central part of Honshu, where blade technology emerged relatively early inJapan. In this region, cultural elements that have been considered part of “Modern Human behavior” appeared
gradually, indicating that the Upper Palaeolithic cultures did not spread out at once. Lastly, it is important in
future to examine regionally diverse processes towards the Upper Palaeolithic in the Japanese archipelago as it isgeographically extensive and diverse in the north-south direction.

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On the mechanics of the spear thrower

Yasuo Higurashi

Graduate School of Human Sciences, Osaka University, Japan. Email: higurasi@hus.osaka-u.ac.jp

Introduction
Between about 50,000 and 40,000 years ago, Neanderthals in Europe were replaced by anatomically Modern
Humans. Various explanations have been offered for processes leading to the replacement of Neanderthals byModern Humans (RNMH). One argument has posited that Modern Humans developed complex projectile
weapons, such as the spear thrower, before the RNMH, and that these innovative technologies improved hunting
efficiency and broadened the range of prey species availability. (The spear thrower is a tool that increases theeffectiveness of spear throwing, and consists of a shaft with a hook or socket at the end that propels the tail of thespear.) To properly evaluate this argument, empirical data are needed on spear throwing with a spear thrower.
In the project entitled “Reconstruction of Hunting Behavior with Projectile Weapons in Neanderthals and Early
Modern Humans Based on Differences in Skeletal Morphology”, I have been making experimental studies 1)
to show the respective advantages of using the hand-cast spear and spear thrower in hunting and 2) to estimate
spear-throwing ability in Neanderthals and early Modern Humans. In this meeting, I report the results of two
experiments concerning the mechanics of the spear thrower.

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Experiment 1
The purpose of Experiment 1 was to investigate similarities and differences in upper-limb kinematics between
spear throwing with a spear thrower and by hand alone (Fig. 1). Two healthy subjects with no special experiencein throwing spear-like objects participated in this experiment. The experiment was conducted outdoors at Osaka
University. The subjects were instructed both to throw a spear as quickly as possible and to aim at a stationary
circular target (0.8 m diameter) placed 8 m away. Throws were videotaped using three digital camcorders at
a sampling rate of 240 Hz. The bony landmarks and proximal end of the spear were digitized using FrameDIAS IV motion analysis software, and their three-dimensional coordinates were obtained using the direct
linear transformation technique. The velocities of the digitized points and the joint angles were compared
between the two throwing methods. The experiment quantitatively showed similarities and differences in upperlimb kinematics between spear throwing with a spear thrower and by hand alone, and reveals an aspect of themechanics of the spear thrower.
Experiment 2
The purpose of Experiment 2 was to examine the kinetic energy transfer from the body to the spear. Ten healthy
subjects with no special experience in throwing spear-like objects participated in this experiment, and theexperiment was conducted outdoors at Osaka University. The subjects performed six different tasks: 1) throwing
spears horizontally with a spear thrower; 2) throwing spears horizontally by hand alone; 3) throwing spears
obliquely with a spear thrower; 4) throwing spears obliquely by hand alone; 5) throwing spears with a spear
thrower while sitting; and 6) throwing spears by hand alone while sitting. Spear and body movements during
throwing were analyzed using a video-based motion-capture system (Frame-DIAS IV ). The results showed

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that the use of a spear thrower generally increased the spear release velocity and the kinetic energy transfer fromthe body to the spear. How the characteristics of the spear thrower shown in this study work in a real hunting
situation are discussed.

Fig. 1 Spear throwing with a spear thrower (a) and by hand alone (b).

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Mediated action through art works among children:
Similarities and differences between traditional and modern
society
Eiko Yamagami

Kobe Gakuin University, Japan. Email: yamagami@human.kobegakuin.ac.jp

The concept of mediated action was originally advocated by Vygotsky (1934) from a sociocultural perspective.
That is, he insisted that there were three basic themes: 1) reliance on genetic and or developmental analysis,2) the claim that higher mental functioning in the individual derives from social life, 3) the claim that humanaction, on both the social and individual aspects, is mediated by tools and signs. Although these three themes areclosely intertwined in his theory, in this study, I will focus on the third one. Mediated action carried out by meansof psychological tools (signs), such as languages, maps, calculation system, art and so on, is peculiar to humanbeings. In particular, art works, which were chosen as media here, involve both concrete materials such as paintand canvas in the outer world and psychological meaning such as desire and phantasy in the inner world. That is,
art is one of the “transitional objects” (Winnicott 1973), which functions as a bridge between reality and mind,
and that eventually can be a representation with the effect of catharsis and sublimation (Freud 1915) and symbols
with transcendent function ( Jung 1928).
The purpose of this study is exploring mediated action among children through art works, which might
demonstrate that human beings unique and effective in responding to the environment. I also find there is adifference between traditional and modern society in terms of mediated action. As for the procedure, I applied
drawing and collage to Baka Pygmy children in Cameroon, as well as to modern society children in Japan andthe USA. I organized small groups, which were composed of developmental stage such as early childhood (56yr.), middle childhood (10yr.) and adolescence (12yr. over), separate sexes, and age and sex mixed. Each groupconsisted of 2-6 members. The instruction was “draw what you ate yesterday”, “draw yourself at that time” and “drawthe person you ate with”. The collage was administered with the instruction of “regard this paper as your house,
and then cut and paste what you want here in your house out of these photos”.

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The results supported the previous research, in which “socialization practices in some nonwestern cultures
involve much less reliance on verbal communication than is typical for western children” (Rogoff 1990). Instead,nonwestern children had visual spatial memory tasks involving eyesight and hearing. Actually, in this study
Baka children did not talk so much but were sensitive and responded to each other, using pointing, gazing andmurmuring. This nonverbal (preverbal) communication was observed for them. For example, in the collage work,
the children were required to use scissors and glue. One of them was not able to use scissors and looked up toan older boy. Then the older one understood the implication from his facial expression and cut materials insteadof him without words (Fig. 1). On the contrary, in the group of 5 years old boys in Japan, one boy asked anothermember how to paste, and he responded with words, helping him. On the drawing, the media of sheet of paperand crayon might facilitate Baka children to express their fun time with a football, important legs for forest life
as well as wild food. Although these members did not discuss the drawing with each other, all of the exposed
images were similar (Fig. 2). On the other hand, in a young group in Japan, two of them drew “spaghetti, I, mom,dad and sister”, while talking to decide what they should draw (Fig. 3). Thus, although there was difference ofcommunication style between traditional and modern society, the media can facilitate children to cooperate withothers.
In conclusion, cooperation among children was observed through the mediated action, such as “an innate,

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Poster Presentationsspecies-specific capacity for shared intentionality” (Tomasello 2009), and moreover, “an innate, species-specificcombinational capacity” (Spelke 2009).
References
Freu S (1915) Zeitschrift fur Psychoanalyse. (Imura T trans. (1970) Freud-Theory of Ego. Nihonkyobunsha, Tokyo.)Jung CG (1928) Die Beziehunggen zwischen dem Ich und dem Unbewuβten. Reichl, Darmstadt. (Matsushiro Y, Watanabe M
trans. (1995) Ego and Unconsciousness. Daisanbunmeisha, Tokyo.)Rogoff B (1990) Apprenticeship in thinking: Cognitive development in social context. Oxford University Press, New York.
Spelke ES (2009) Forum. In: Tomasello M, Why we cooperate. The MIT Press, A Boston Review Book, Cambridge, Mass andLondon, pp. 149-172.
Tomasello M (2009) Why we cooperate. The MIT Press, A Boston Review Book, Cambridge, Mass and London.
Vygotsky LS (1934) Myshlenie i rech’: Psikhologicheskie issledovaniya. [Thinking and speech: Psychological investigations.]
Sotsial’no-Ekonomicheskoe Izdatel’stvo, Gosudarstvennoe, Moscow and Leningrad.
Wertsch JW (1991) Voices of the mind. A sociocultural approach to mediated action. Harvard University Press, Cambridge,
Mass.
Winnicott DW (1971) Playing and reality. Tavistock Publications, London.

Fig. 1 Baka boy’s group: Football, vehicles, animals.

Fig. 3 Japanese 5 yr. children: Spaghetti, I, mom,dad, sister, grapes, salads.

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Fig. 2 Baka girl’s group: Humans with exaggerated
legs, footballs, spears, fishes, chickens, potatos.

Cognitive development of Baka Pygmy children growing up
in modern hunter-gatherer society
Tadashi Koyama

Faculty of Humanities and Sciences, Kobe Gakuin University, Japan. Email : t-koyama@human.kobegakuin.ac.jp

Cognitive development of the children in growing up in hunter-gatherer society
Homo sapiens had to adapt to their environment. To do that, we have developed more complex knowledge andbehavior. Development of more complex knowledge and behavior is dependent on dynamic interplay among
biological, cognitive, and social system dimensions (Davis and Bedore 2013). For example, in language learning,
infants would do differentiate between salient and non-essential incoming stimuli, gain and store more diverse
goal-directed movements related to their ambient language requirement by infant-internal interconnections
between physical systems of body and social interaction capacities tuned by input of ambient language (Davis andBedore 2013).
The children who are grown up in hunter-gatherer society would develop their cognitive capacity to continue
to maintain the life of their society. In this paper, the author investigated the distinctiveness of cognitive
development in Baka children who growing up in modern hunter-gatherer society.
Repetitive production

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In Fig. 1 overlapping circles indicate physical capacities involved in acquisition of knowledge structures. For
example, that forms the basis of fully realized phonological system (Davis and Bedore 2013). Each relies on an
overall general capability for pattern formation. Pattern formation would be tuned and refined (it is the cognitive
process) by extrinsic context in early infancy by development of joint attention, intention reading, and turn taking.
Refining patterns affect the productions.
Homo sapiens adapted to their environment by tuning and refining with cognition and repetitive production.
Neanderthals also might have used this process. The capacity of imagination might develop on the base of tuning,refining and repetitive production. Those processes would be noted for early human adaptations.
The process of tuning and refining would affect sharing intentionality
The capacity of infants is to be required for their future living. In Baka infants who are grown up in modern
hunter-gatherer society, about 9 month or so, infant would begin to tune to ambient inputs and begin toincorporate into them (Koyama 2014). Maybe joint attentional behaviors led to activate the tuning process. It
was reflected in children’s social referencing behavior indicating sharing intentionality of others.
The author investigated social referencing behaviors of 27 Baka children during performing object-construction
with wooden blocks in accordance with model which a stranger made. Their parents permitted to theparticipation in this study. Children were introduced to the play situation with the author by an assistant who
was a Japanese anthropologist being acquainted with Baka people participating this study. He could speak Bakalanguage well. And he was there in the situations a child played with the author. During that play children
engaged in constructing object with wooden blocks matching with a model made by the author in the face to face

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situation. The verbal instructions to the children were translated into Baka language.
Fig. 2 indicated the mean frequencies of children’s social referencing to the author in each age group. Table 1
showed SD of each age group. It was thought that results showed sharing with intentionality of others during
construction with brocks in accordance with model that adults made. At just about 4 years of age, socialreferencing to author during children’s constructing (in process) and performing goal directed acts (completed)checking form the viewpoint of others increased. Those behaviors showed sharing intentionality with others inconstruction of objects. We need to investigate the relationship between these behaviors and children’s flexible
adaptation from about 4-year-old.
Their capacity of abstraction
As I reported in previously, Baka children showed much flexibility on object substitution in their play andmatching up geometric figures (Koyama 2013, 2014). Matching up geometric figures indicated their capacity of
abstraction of shapes. As for those developments, variation was found (Koyama 2014). The capacity of abstraction
would develop gradually from 3 year-olds to 5 years-olds (Table 1).
Entering into languages in figures: Baka children were in oral language culture
Nelson (2014) pointed that in cultural evolution there were three stages in human development. The first one ismimetic culture stage. Chimpanzees might be in this stage. Next is oral language culture stage. Nelson (2014)
emphasized narratives were main mythology for children. Third is the stage of language in figures. Baka children
were in oral language culture, and they began to enter into language in figures culture and they showed intuition
to perception of figures (Koyama 2012). They could collect and match one picture with a similar figure that was
depicted objects they had not seen and gone through that. But using figures in their symbolic play were seldom
observed in my research.
Distinctiveness in their pretend play
The copy of daily experiences is noticed in Baka children’s play. The author hypothesizes that human imagination
develops on the base of anchoring to their real world and reconstructing daily experiences and our knowledge(decoupling) (Leslie 1987; Koyama 2013). Those processes were observed in Baka children’s pretended play.
Baka children shared object substitutions with other children (Koyama 2013). Production with objects in theirsymbolic play based on copying of their experiences and older people’s daily activities in daily hunter-gathering
life was observed. That plays were cooperative with other children. From about two years old, they simulated older
people’s daily activities in those play. Those simulations were noticed. In addition to that, tuning and refining
based on previous repetitive production and objects pragmatism were showed in those plays. And this would be
noted from the view point of embodied cognition.

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References
Davis BL, Bedore LM (2013) An emergence approach to speech acquisition: Doing and knowing. Psychology Press, London.
Koyama T (2012) Comprehension of pictures in Baka children who grow up in the hunter gatherer culture. In: Terashima H (ed)Replacement of Neanderthals by Modern Humans: A-02 Team Annual Report 2. Kobe Gakuin University, Kobe, pp. 55-60. (InJapanese)
Koyama T (2013) Cognitive flexibility and making objects in Baka Pygmy children. In: Akazawa T, Ogihara N, Tanabe HC,
Terashima H (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 2: Cognitive and physical perspectives.
Springer, New York, pp. 33-38.
Koyama T (2014) Development of metarepresentation in Baka Pygmy children. In: Terashima H (ed) Replacement of Neanderthals by Modern Humans: A-02 Team Annual Report 4. Kobe Gakuin University, Kobe, pp. 41-44. (In Japanese)

Poster Presentations

Koyama T (2014) Development in geometric figures-recognition of Baka Pygmy children. 44th Annual Meeting of the Jean
Piaget Society, San Francisco, Poster Session 4, 30 May 2014.
Leslie AM (1987) Pretence and representation: Origins of “thory of mind.” Psychological Review 94: 412-426.
Koyama T (2014) Stability and flexibility in early social cognitive development (in submission).
Nelson K (2014) The cultural basis of language and cognitive development. 44th Annual Meeting of the Jean Piaget Society, San
Francisco, Plenary 3, 29-31 May 2014.

Fig. 1 Physical capacities involved in acquisition ofknowledge structures.
Fig. 2 Mean of frequencies of social referencing in Baka
children while they were engaged in constructing object withwooden blocks.

Table 1 SD of each age group.

3y
4y
5y

in process completeds
1.03
1.48
4.47
1.2
0.79
0.49

Table 2 Percentage of choosing same figure in type I (Koyama 2014).

3 year olds
4 year olds
5 year olds

circle
30.0
50.0
85.7

triangle
50.0
70.0
71.4

square
50.0
70.0
71.4

semicircular
50.0
70.0
71.4

cross
60.0
80.0
71.4

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Learning process on the skills of reading nature

Kaoru Imamura1* and Hiroyuki Akiyama2

1. Nagoya Gakuin University, Japan; 2. Kyoto Kacho University, Japan. *Email: kaoru.imamura@gmail.com

The foraging strategy of the Neanderthals is supposed to have centered around large mammals. On the other
hand, the tool manufacturing industry of early Modern Humans have shown a clear trend to the diversificationof food items (Fa et al. 2013; Kuhn and Stiner 2006; Richards and Trinkaus 2009; Snodgrass and Leonard2009), which shows the development of cognitive capacity on their envioronment and food resources, i.e., nature.
The innovation of the skills of ‘reading nature’ might have occurred in Modern Humans (Shipman 2011). To
understand the characteristics of learning the skills of reading nature, this study focuses on learning process ofhunting skills among children of the San, hunter-gatherers in the Kalahari Desert of southern Africa.

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San are well-known as large game hunters shooting with a bow and arrow, and/or a spear. However, they get meat
animals, not only large but also middle and small size animals and birds, by various methods, especially by snares.
Traps provide a stable supply of animal meat. I collected information about traps. The results are;
1. They have many kinds of traps for large game such as gemsbok and kudu, middle size mammals such as duiker
and steenbok, small mammal such as mongoose and ground squirrel, middle and large size birds such as ostrich,kori bustard and korhaan, and small birds.
2. Not only men but also women use snares especially for middle and large sized birds.
3. Boys play with trapping small animals.
Focusing on boys hunting, they use snares for small animals and middle size birds, a toy bow and arrow for birds
and lizards, a slingshot for small animals, birds and lizards, and traps with waylay for catching small birds. On theother side, girls learn how to set snares for birds when they go gathering with women. To get technic for traps
children must know the behavior of animal and which vegetables animal eat. They learn ‘skills of reading nature’by hunting.
References
Fa JE, Stewart JR, Lloveras L, Vargas JM (2013) Rabbits and hominin survival in Iberia. Journal of Human Evolution 64: 233241.
Kuhn SL, Stiner MC (2006) What’s a mother to do? The division of labor among Neandertals and Modern Human in Eurasia.
Current Anthropology 47: 953-980.
Richards MP, Trinkaus E (2009) Isotopic evidence for the diets of European Neanderthals and early Modern Humans. Proceedings of the National Academy of Sciences 106: 16034-16039.
Shipman P (2011) The animal connection: A new perspective on what makes us human. Norton, New York.
Snodgrass JJ, Leonard WR (2009) Neanderthal energetics revisited: Insights into population dynamics and life history evolution.
PaleoAnthropology 2009: 220-237.

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Fertility and population dynamics of Baka hunter-gatherers
in southeast Cameroon
Izumi Hagino1* and Taro Yamauchi2

1. Graduate School of Health Sciences, Hokkaido University, Japan; 2. Faculty of Health Sciences, Hokkaido University, Japan.
*Email: hagino.is.me@gmail.com

Cognitive development of the children in growing up in hunter-gatherer society
Demographic parameters adequately indicate the health status of a small population. Until the 1980s, studies of
Pygmy populations described low viability and very short life expectancies (mid-20s). The Baka are a group of
pygmy hunter-gatherers living in the central African rainforest whose habitat extends mainly over southeastern
Cameroon. In the last few decades, Baka society has shifted to settlements; communities relying entirely on
hunting and gathering are disappearing. Permanent villages along roads increase access to public health centers.
There now exists intergenerational variation among their demographic characteristics. This study aimed, namely,
to reveal the fertility and population dynamics of current Baka hunter-gatherers and, to estimate the transition of
their fertility over the generations.
Subjects and methods
Census, birth, death, and migration data for four Baka villages located in the East Province of Cameroon werecollected three times from February to March of 2011, 2012, and 2013. Personal interviews covering names, clans,
marital statuses, and births of children were conducted with all residents. Their ages were estimated in one-year
ranges for children and five-year ranges for adults. Genealogical data was collected from 66 women who married
village residents. There were a total of 295 women who had completed reproduction in reconstituted pedigree.
Following Ohtsuka (1990), they were categorized into four generational groups: (A) not all female offspring
have married; (B) All female offspring have married; (C) all female offspring have completed reproduction; (D)All female offspring of all female offspring have married. The daughter-mother ratio, (DMR), the ratio of thenumber of married female children to the number of mothers, was calculated for each group.

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Results and discussion
Fig. 1 shows the population structure in 2011; the population was 611 and the sex ratio was 107.8. The medianage was estimated at 19.1 years, and 42.6% of population was under 15 years indicating that the Baka population
was relatively young. The number of births exceeded the number of deaths during the observation periods; thus,
the population naturally increased. The crude birth rates were 39.2 for 2011-2012 and 42.5 for 2012-2013. Fig.
2 shows the distribution of the 295 women by number of offspring. The mean completed parity was 5.03 ±3.05. Baka population had the same level of fertility as other African hunter-gatherers as seen in Table 1. Table
2 contains DMR and estimated Lotka rate (P) for each generational group. Although there is no remarkabledifference between the generational groups, nor an increasing trend, the DMR of the this population is relatively
high.
References
Ohtsuka R (1990) Population growth in the past. In: Ohtsuka R, Suzuki T (eds) Population ecology of human survival. University of Tokyo Press, Tokyo.

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Fig. 1 Population structure of target villages in 2011.

Fig. 2 Percent distribution of number of children of both sexes.

Table 1 The fertility among hunter-gatherer societies.

group
Baka
Aka
!Kung
Agta
Ache

Fertility
5.03
5.04
4.69
6.53
8.03

Study area
Cameroon, rainforest
Central African Republic, rainforest
Namibia, Kalahari desert
Philippine, rainforest
Paraguay, rainforest

Reference
present study
Hewlett, 1978
Howell, 1979
Goodman et al., 1985
Hill and Hurtado, 1996

Table 2 Daughter-Mother Ratio (DMR) and estimated Lotka rate (P) by mother's age groups.

group
B
C
D
B-D

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N
88
37
26
151

No. of married children
Male
Female
106
235
62
83
38
81
206
399

DMR
2.670
2.243
3.115
2.642

P at mean child bearing age
20
25
0.050
0.040
0.041
0.033
0.058
0.047
0.050
0.040

The formation of cooperative society initiates the evolutionof risk-taking innovative behaviors
Ryosuke Kimura1*, Wataru Nakahashi2 and Kohei Tamura3,4

1. Graduate School of Medicine, University of the Ryukyus, Japan; 2. School of Advanced Sciences, Graduate University for Advanced
Studies, Japan; 3. Graduate School of Information Science and Technology, The University of Tokyo, Japan; 4. CREST, Japan.
*Email: rkimura@med.u-ryukyu.ac.jp

Introduction
Modern Humans are outstanding in innovative behaviors among all living animals. How did highly innovative
behaviors evolved only in humans? In other words, why did the other primates not acquire such behaviors? Thisis one of the essential questions for understanding the evolution of human learning strategy and ability. Many
researchers have theoretically studied cultural evolution to clarify what factors lead highly developed cultures inhumans (Reviewed in Aoki and Feldman 2014). These previous theoretical studies have pointed out a paradox
that, in the presence of imitation (social learning) strategy, risk-taking innovation behaviors do not survive andthus cultural evolution is not promoted. Therefore, some model will be required to break such a paradox. In
this study, we theoretically examined the possibility that the human-specific cooperative society can initiate theevolution of innovative behaviors.
Methods
We computationally simulate a co-evolutionary model of cooperation and learning strategies. The schematic
representation of the model is shown in Fig. 1. In this model, we assumed two genetically determined behavioral
traits: “individualism/collectivism” and “individual learning (IL)/social learning (SL)”. Each generation
experiences a lifecycle including (1) birth, (2) mutations, (3) innovations by IL individuals, (4) horizontal learning
by SL individuals, (5) obtaining resources, and (6) sharing resources by collectivists. Initially, all individuals haveindividualism and IL alleles, and the simulation runs 10,000 generations. Key parameters in this simulation are
the benefit (cost) of collectivism (sharing resources), α, and the number of groups, L, that reflects the number ofindividuals in each group. We observed the number of individuals that genetically adapt each strategy. Then, thespeed of cultural evolution was compared between various conditions. To confirm the results of the simulation,mathematical analyses were also performed.

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Results
When collectivism is not beneficial (α ≦ 0), collectivists cannot survive in the population (Figs. 2a and b). In
addition, SL individuals almost occupy the population under this condition. This indicates that IL individuals
are weaker than SL individuals under the competition between individuals in our simulation. As collectivism
becomes beneficial (α>0), however, the number of IL individuals increases (Figs. 2c and d), suggesting thatcollectivism rescues the survival of IL individuals. The number of groups, which reflects the number of individuals
belonging to each group, also affects the survival of IL individuals (Figs. 2e and f ). The number of rescued IL
individuals increases with increasing the number of individuals in each sharing group. The speed of cultural
evolution depends on the number of IL individuals in the population (Fig. 3).
Discussion
Using computer simulations, the present study suggested that a resource-sharing society rescues risk-taking

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innovators. In addition, the simulations demonstrated that, as the number of individuals in each sharing group
becomes large, the proportion of innovators and the rate of cultural evolution in the total population increase.
This phenomenon is possibly explained as follows: (1) a sharing group having a comparatively large proportionof IL individuals earns a relatively small total amount of resources in the group, (2) a small number of individuals
in each group results in a large variance in the number of IL individuals among groups, and therefore (3) the
negative pressure on IL individuals becomes stronger as the size of sharing groups is smaller. These results werealso showed by a mathematical analysis under some simplifying assumptions. Our findings indicate the possibility
that formation of cooperative societies, rescuing risk-taking innovators, is associated with the evolution of thehuman learning strategy and ability. Along with learning ability, cooperation is also an important behaviour for
the understanding of human evolution. Our present study is intriguing in the point of connecting these two
human-specific behaviors. It has been suggested that the Neanderthals had small-scale cooperative societies.
Therefore, the size of sharing groups may be one of the factors that sealed the fates of two species, Neanderthalsand Homo sapiens. The “cooperative society-initiated” hypothesis suggests that most of the members in thesociety are social learners and only a small number of innovators sustain the cultural evolution. Future studies willaddress how learning strategy and ability-related genes evolve in such a condition.
References
Aoki K, Feldman MW (2014) Evolution of learning strategies in temporally and spatially variable environments: A review oftheory. Theoretical Population Biology 91: 3-19.

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Fig. 1 Simulation model. (a) Hereditary system. A haploid organism with sexual reproduction is assumed. Genes for learning
strategies (IL/SL) and cooperation strategies (individualism/collectivism) inherit independently from either of parents. The
offspring takes over the technology level (q) from a parent (vertical transmission). (b) Mutations. Genes can reciprocally mutate
at a certain rate (μ = 0.001). (c) Innovations. IL individuals make an innovation. The technological advance (s) of the innovationis stochastically given depending on a function f(s; β) = exp(-s/β)/β – β. (d) Horizontal transmission. SL individuals observe kindividuals from the same generation in the whole population and learn the best technology of them. (e) Sharing resources.
Individuals obtain resources in proportion to their technology. Collectivists share the resources within the group that depart andgather in every generation. Sharing resources yield a benefit (α>0) or cost (α<0).

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Fig. 2 The number of each strategy in the population. The results of 10 simulations were superposed (one in orange and therest in gray). Ind, individualist; Col, collectivist; α, benefit of collectivism; L, the number of groups.

Fig. 3 Cultural evolution. α, benefit of collectivism; L, the number of groups.

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Genome-wide association analysis of cephalic form inmodern Japanese
Kyoko Yamaguchi1*, Toshiyuki Hayakawa2, Chiaki Watanabe1,
Takehiro Sato1, Akira Kawaguchi1, Ken Yamamoto3, Hajime Ishida1 and
Ryosuke Kimura1
1. University of the Ryukyus, Japan; 2. Kyushu University, Japan; 3. School of Medicine, Kurume University, Japan.
*Email: kyoko777@gmail.com

Introduction
Human head size is correlated with the brain size especially during infancy. Wickett et al. (2000) reported asignificant correlation between head perimeter and total brain volume obtained by MRI among healthy adults(r=0.659, P<0.001). Head circumferences is often used as a proxy index for normal brain development in infants,
especially in terms of early detection of autism spectrum disorder (ASD) (Kanner 1968; Raznahan et al. 2013).
The cephalic index (CI), a ratio of the maximum width to the maximum length of a head multiplied by 100, wastraditionally used for clustering human populations in different regions although a few studies have revealed thesignificance of the environmental factors on CI (Kouchi 2004).

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Genetic basis of normal variation in human head form has remained unclear for a long time except for several
genes that are responsible for pathologically abnormal head size, such as MCPH1 for microcephaly. MCPH1
has played an important role in the evolution of Homo sapiens after the divergence from Neanderthals (Greenet al. 2010). Recent genome-wide association studies (GWASs) have begun to find genomic regions associatedwith head size such as infant head circumference (Taal et al. 2012) or adult intracranial volume (Ikram et al.
2012). However, these studies are based on European descendents and may not represent Modern Humans as awhole. In this study, we collected cephalic form data in Japanese and conducted a GWAS to identify the genes
accounting for head form, to reveal the genetic basis of head form among Modern Humans.
Materials and methods
We collected DNA sample as well as the size measurements of head for a total of 767 Japanese people residing
in Okinawa Island, Okinawa, Japan. Genomic DNA was extracted from blood or saliva of participants andgenotyped for 729,172 SNPs using HumanOmniExpress BeadChip system (Illumina, San Diego, CA, USA),and quality control was carried out. The head measurements included the head circumference measured with atape measure, the maximum head length and breadth measured with an anthropometer, and the cephalic index
calculated from the crude measurements. With these head measurements and an index as dependent variables,
GWA analyses were conducted using a whole genome association analysis toolset, PLINK, controlling for effectof sex, height, BMI, and principal components of genotype data. The genes that showed significant associations
with head size and form were then further examined in terms of comparison with Neanderthal and Denisovan
genomes.
Results and discussion

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A single nucleotide polymorphism (SNP), rs10470149 located within LOC284825 on chromosome 21q21.3
was associated with head circumference at a conventional genome-wide significance level (P<5×10-8)(Fig. 1). The
derived allele of this SNP was associated with a smaller head circumference. Among the SNPs that had P-values

Poster Presentations

below a threshold for suggestive association (P<1.0x10-5), rs10236933 fell in CNTNAP2, a gene whose relation to
ASD and schizophrenia has been reported (Rodenas-Cuadrado et al. 2014) (Table 1, Fig. 1).
Our analysis did not find significant association between head circumference and the SNPs that were associatedwith infant head circumference (Taal et al. 2012) or adult intracranial size (Ikram et al. 2012) in previous
GWASs. In these studies, however, the participants are European descendants. Our results suggest different
genetic backgrounds of head size may exist for Asians and Europeans.
As for cephalic index, no SNP showed a significant association at a conventional genome-wide significant level.
This is consistent with the findings that a ratio of head breadth to length is affected by environment whereas asum of breadth and length may be genetically determined (Martínez-Abadías et al. 2009). However, further study
with large sample size is needed to confirm the result of present study.
References
Green RE, Krause J, Briggs AW, Maricic T, Stenzel U, Kircher M, Patterson N, Li H, Zhai WW, Fritz MHY et al (2010) A
Draft Sequence of the Neandertal Genome. Science 328: 710-722.
Ikram MA, Fornage M, Smith AV, Seshadri S, Schmidt R, Debette S, Vrooman HA, Sigurdsson S, Ropele S, Taal HR et al (2012)
Common variants at 6q22 and 17q21 are associated with intracranial volume. Nature Genetics 44: 539-544.
Kanner L (1968) Autistic disturbances of affective contact. Acta Paedopsychiatr 35: 100-136.
Kouchi M (2004) Secular changes in the Japanese head form viewed from somatometric data. Anthropological Science 112: 4152.
Martínez-Abadías N, Esparza M, Sjøvold T, González-José R, Santos M, Hernández M (2009) Heritability of human cranial
dimensions: comparing the evolvability of different cranial regions. Journal of Anatomy 214: 19-35.
Raznahan A, Wallace GL, Antezana L, Greenstein D, Lenroot R, Thurm A, Gozzi M, Spence S, Martin A, Swedo SE et al (2013)
Compared to what? Early brain overgrowth in autism and the perils of population norms. Biological Psychiatry 74: 563-575.
Rodenas-Cuadrado P, Ho J, Vernes SC (2014) Shining a light on CNTNAP2: Complex functions to complex disorders. European Journal of Human Genetics 22: 171-178.
Taal HR, St Pourcain B, Thiering E, Das S, Mook-Kanamori DO, Warrington NM, Kaakinen M, Kreiner-Moller E, Bradfield
JP, Freathy RM et al (2012) Common variants at 12q15 and 12q24 are associated with infant head circumference. Nature Genetics 44: 532-538.
Wickett JC, Vernon PA, Lee DH (2000) Relationships between factors of intelligence and brain volume. Personality and Individual Differences 29: 1095-1122.

Fig. 1 A Manhattan plot of a genomewide association analysis for headcircumference. Each dot representsa P-value of a SNP for an association
after controlling for sex, height, BMI, andpopulation structure (principal components
1 and 2 of genomic variance).

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Table 1 A list of SNPs that had P-values below the suggestive level (1.0x10-5) for associations with head circumference. Linear
regression analyses were conducted controlling for sex, height, BMI, and population structure (principal components 1 and 2 ofgenomic variance).

CHR
BP
21
29468403
1862085800
1616052973
18620875121616053729
7 146337383
1 111908713
7 146356208
820102006

SNP
rs10470149
rs2959456
rs215101
rs2909659
rs4148333
rs10236933
rs1361049
rs2533089
rs2173115

675342884

rs2485067

16

16054694

rs215099

Gene
Derived DAF
LOC284825
C
0.1812
LOC284294
G
0.5945
ABCC1
G* 0.3129
LOC284294
A
0.5932
ABCC1
T
0.6812
CNTNAP2
C
0.2888(PIF, PGCP1)
C
0.2529CNTNAP2G
0.3005(LZTS1, ATP6V1B2) G
0.5391
RP11-554D15.1
T
0.1102(LOC101928516)
ABCC1
A
0.3136Beta-0.4511-0.3238-0.3368-0.31130.321-0.3247-0.3214-0.3094-0.2814P
2.08E-08
4.67E-07
9.70E-07
1.26E-06
2.98E-06
3.07E-06
7.70E-06
7.87E-06
8.27E-06

P(GC) BONF
2.11E-08 0.04353
4.73E-07 0.9777
9.81E-07
1
1.27E-06
1
3.01E-06
1
3.10E-06
1
7.78E-06
1
7.94E-06
1
8.35E-06
1-0.4457

8.63E-06

8.71E-06

1-0.3068

9.13E-06

9.22E-06

1

CHR: Chromosome, BP: Base pair position on a chromosome based on GRCH37/hg19, SNP: Reference SNP ID number, DAF: Derived allele
frequency, Beta: Regression coefficient of the derived allele for the association with head circumference after controlling for sex, height, BMI andgenomic PC1and PC2, P: crude P-value for the association without genomic control, P(GC): P-value after genomic control, BONF: P-value after
Bonferroni correction for the number of SNPs.
*: Minor allele instead of derived allele.

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The effect of trauma on cultural transmission of Neanderthals

Wataru Nakahashi

The Graduate University for Advanced Studies, Japan. Email: w.nakahashi@gmail.com

In many cases, learning how to use our body is necessary to master difficult cultural skills. More advanced cultural
skills tend to be more complex and demand longer training of body use. We have to memorize cultural skills not
only in our brain but also in our body. Therefore, when an individual suffers a serious traumatic injury on a bodypart associated with a cultural skill, thereafter he/she may not hold the skill because of the deteriorated physicalability. Since the medical technology of archaic human society was very poor, traumatic injuries may have strongly
affected physical ability of individuals, which may have restricted the advancement of the culture.
An analysis of traumatic lesions among Neanderthals revealed that they frequently suffered traumatic injuries on
upper body (Berger and Trinkaus 1995). Since upper body is the most important part for humans to hold cultural
skills, we can suppose that high injury rate inhibited the maintenance of difficult cultural skills in Neanderthalsociety. However, how their injury affected their culture has not well investigated, although what kind of culturecaused their injuries had been discussed (Berger and Trinkaus 1995; Trinkaus 2012).
To maintain an advanced cultural skill (which can be acquired only by social learning) in the population, an“expert” of the skill must successfully transmit the skill to more than one “novice” of the skill before he/she diesor injures to lose the skill. In other words, before one expert disappears, more than one expert must emerge inthe population. When injury rate is high, the skill is less likely to be maintained in the population because two
possibilities increase; a possibility that an expert injures and lose the skill before he/she transmits it to novices,
and a possibility that a novice already injured and lose the ability for mastering the skill before he/she has acommunication with an expert. Therefore, when we take the effect of injury into consideration, each individual
must transmit his/her cultural skill to more than one individuals during his/her lifetime. The necessary numberof transmittable individuals for the maintenance of a cultural skill in the population depends on the injury rate
associated with the cultural skill. In this study we consider the transmission of a series of cultural skills associatedwith small animal hunting by complex projectile weapons, which may have been absent in Neanderthal society
and appeared after the emergence of Modern Humans (Shea and Sisk 2010).
To obtain the necessary number of transmittable individuals corresponding to the injury rate of Neanderthals,in this study, we consider the following cultural transmission model, which is described in Fig. 1. This modelis an arrangement of the SIR model that is often used in mathematical biology to describe a transmission ofcommunicable disease through individuals. We consider cultural transmission of a complex cultural skill which
can be acquired only by adults. In the adult population, there are three kinds of individuals, novices (N) who haveneither the cultural skill nor serious trauma, experts (E) who have the skill without trauma, and traumatizedindividuals (T) who suffered serious traumatic injury and lose the ability for holding the skill. When a novice
closely communicates with an expert, he/she sometimes acquires the advanced skill and becomes an expert. The
rate of cultural transmission is β. Both novices and experts sometimes suffers a serious traumatic injury and lose
the ability for holding the advanced skill. Traumatized individuals never revert to be novices and experts. The
injury rate is γ . Every individual has a same death rate, μ , regardless of the kind of the individual, but new adults
emerge in the population with the same rate, μ , so that population size is always constant. There is a possibility
that an individual injures before adulthood. The proportion of non-injured individuals in new adults is h. Writing

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the changes of frequencies of three kinds of individuals by differential equations, we have
(1a)
(1b)
(1c)
Here, N, E, and T are the proportions of novices, experts, and traumatized individuals in the adult population,
respectively, so that always holds.
We include Neanderthal fossil data in the above model to estimate parameters of Neanderthals. From Trinkaus(2011) and Caspari and Lee (2006), the annual death rate of adult Neanderthals, μ , isestimated around 0.070.085. From Berger and Trinkaus (1995), Wu et al. (2011), and the Catalogue of Fossil Hominids Database,the proportion of traumatized individuals who lost the capacity for cultural skills associated with small animal
hunting by complex projectile weapon, T̂ , is estimated around 0.5-0.6. From Trinkaus et al. (1994), Berger andTrinkaus (1995) and Trinkaus (2011), the proportion of non-injured individuals in new adults, h, is estimatedaround 0.8-0.85. Including these values into the mathematical model, we can estimate the annual injury rateof adult Neanderthals, γ , the critical cultural transmission rate of Neanderthals, β* , and the critical number ofcommunicable individuals during adulthood in Neanderthals, β* /μ , shown in Table 1.
Note that to obtain the critical number above, we assume that the cultural skill is always transmitted froman expert to a novice when they closely communicate with each other. However, in reality, cultural traits are
not necessarily transmitted between them even when they have a close communication, because some experts
learn wrongly, do not have a motivation to learn, or some experts do not demonstrate the skill. Therefore,
communicating far more than 4-6 individuals, probably 10-15 individuals, during adulthood may have been
actually necessary for maintaining the cultural skills in Neanderthal society.
Ancient genome sequences suggest that Neanderthals may have commonly mated among close relatives probably
because of small community size (Prüfer et al. 2014; Castellano et al. 2014). Therefore, it may have been almost
impossible for Neanderthals to communicate closely with more than 10-15 individuals during adulthood. Thus,
we can conclude that it was difficult for Neanderthals to maintain advanced cultural skills that demand a masteryof difficult physical techniques, because their injury rate was too high and their community size was too small.

152

References
Berger TD, Trinkaus E (1995) Patterns of trauma among the Neandertals. Journal of Archaeological Science 22: 841-852.
Caspari R, Lee SH (2006) Is human longevity a consequence of cultural change or modern biology? American Journal of Physical Anthropology 129: 512-517.
Castellano S, Parra G, Sánchez-Quinto FA, Racimo F, Kuhlwilm M, Kircher M, ... Pääbo S (2014) Patterns of coding variationin the complete exomes of three Neandertals. Proceedings of the National Academy of Sciences 111: 6666-6671.
Prüfer K, Racimo F, Patterson N, Jay F, Sankararaman S, Sawyer S, ... Pääbo S (2014) The complete genome sequence of a Neanderthal from the Altai Mountains. Nature 505: 43-49.
Shea JJ, Sisk ML (2010) Complex projectile technology and Homo sapiens dispersal into western Eurasia. PaleoAnthropology
2010: 100-122.
Trinkaus E (2011) Late Pleistocene adult mortality patterns and Modern Human establishment. Proceedings of the National
Academy of Sciences 108: 1267-1271.
Trinkaus E (2012) Neandertals, early Modern Humans, and rodeo riders. Journal of Archaeological Science 39: 3691-3693.
Trinkaus E, Churchill SE, Ruff CB (1994) Postcranial robusticity in Homo. II: Humeral bilateral asymmetry and bone plasticity.

Poster Presentations

American Journal of Physical Anthropology 93: 1-34.
Wu XJ, Schepartz LA, Liu W, Trinkaus E (2011) Antemortem trauma and survival in the late Middle Pleistocene human cranium from Maba, South China. Proceedings of the National Academy of Sciences 108: 19558-19562.
The Catalogue of Fossil Hominids Database. http://fossil.kochi-tech.ac.jp/

Fig. 1 The description of the NET model.

Table 1 Estimated values of parameters.

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A population-genetics based model for explaining apparent
cultural continuity from the Middle to Upper Palaeolithic in
Eurasia
Yutaka Kobayashi1*, Seiji Kadowaki2 and Masaki Naganuma3
1. Kochi University of Technology, Japan; 2. Nagoya University, Japan; 3. Hokkaido University, Japan.
*Email: kobayashi.yutaka@kochi-tech.ac.jp

Aim
In this study, we construct a mathematical model to simulate the invasion of Homo sapiens (HS) into a region
preoccupied by an archaic hominin (AH) species and associated acculturation on the side of HS. The purpose is
to identify the major determinants of the pattern of cultural shifts associated with biological replacement. We
analyze the model using analytical approximation techniques and individual-based simulations. Based on the
results, we discuss the possible causes of a variation between different regions in the pattern of cultural shifts
during the MP-UP transition.
Model

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Our model is simplistic in many respects to extract essence and also to maintain analytical tractability. Imagine
that a small population of HS is invading into a region preoccupied by an AH species. Suppose that this region
accommodates a total of exactly N individuals of the AH or HS, where N is constant. Suppose, again forsimplicity, that those populations reproduce asexually, so that each individual has only one parent. Generations
are discrete in so far as cultural transmission occurs. HS is initially rare in frequency, given that they have justarrived at this region. Let p denote the frequency of HS in the focal region and p0 the initial frequency. The AH
is less viable than HS for some unspecified reason and therefore the latter increases in frequency and eventuallyreplaces the former. We assume that the relative fitnesses of HS and the AH are 1 and 1-s, respectively. Since thepopulation size is finite, HS may undergo local extinction with a positive probability due to random sampling
drift despite selective superiority. However, we set Ns sufficiently large, so that the probability of extinction for
HS is negligible.
We assume that cultural transmission may occur from the AH to HS, while we neglect that in the opposite
direction for simplicity. We assume that the AH and HS initially have different phenotypes with respect to acultural element, referred to as CA and CH, respectively. We assume that each individual of HS learns verticallyand obliquely with probabilities 1-γ and γ, respectively. When an individual of HS learns obliquely, he/she may
copy either a conspecific or an individual of the AH in the parental generation. As a result, the cultural trait CA
can gradually spread into the population of HS. We assume that the probability that an individual of HS copies
an individual of the AH, denoted by X(u,p), is given byX(u,p) =(u(1-p))/((1-u)p+u(1-p)),

154

(1)

where u represents the tendency to learn from the AH and may range from 0 to 1; u=0 represents an extreme
situation where HS never copies the AH. On the other hand, u=1 implies that individuals of HS, when they learn
obliquely, always choose to copy the AH. When u=1/2, HS has no preference with respect to the role models
for oblique social learning. It must be noted that the probability of learning CA may be higher than equation (1)because individuals of HS may also bear CA. We assume that transmission of a cultural character is always faithful

Poster Presentations

and perfect, so that social learning never fails.
We assume that the two phenotypes CA and CH are selectively neutral. We investigate how the pattern of culturaldynamics depends on selection coefficient s, initial frequency p0, oblique transmission rate γ, and tendency
to learn from the AH u. We are particularly interested in the probability that CA persists despite biologicalreplacement and also the time until it is lost after the replacement.
Results
We can loosely categorize the patterns of cultural dynamics based on some results of individual-based simulations.
Pattern A (Immediate loss of CA): CA is quickly lost during biological replacement. This pattern occurs whenselection is strong (s is large), the initial frequency of HS is high (p0 is large), the preference for the heterospecific culture is low (u is small), and/or the oblique transmission rate is low (γ is small).
Pattern B (CA keeps to be frequent): CA continues to be very frequent during biological replacement and persists forever. This pattern occurs when selection is weak (s is small), the initial frequency of HS is low (p0 issmall), the preference for the heterospecific culture is high (u is large), and/or the oblique transmission rate ishigh (γ is large).
Pattern C (Gradual loss of CA): CA decreases in frequency during biological replacement. It persists for a longtime after the replacement but eventually goes extinct. This pattern occurs under conditions intermediate between the patterns A and B.
Pattern D (Temporal decrease in the frequency of CA): CA reduces its frequency during biological replacement. However, it increases in frequency again, eventually reaches fixation, and persists forever. This patternalso occurs under conditions intermediate between A and B.

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Testing the hypothesis on cognitive evolution of Modern
Humans’ learning ability: Achievements of past-climatic
approaches
Minoru Yoneda1*, Ayako Abe-Ouchi2, Hodaga Kawahata2,
Yusuke Yokoyama2 and Takashi Oguchi3

1. The University Museum, The University of Tokyo, Japan; 2. Atmosphere and Ocean Research Institute, The University of Tokyo,
Japan; 3. Center for Spatial Information Science, The University of Tokyo, Japan. *Email: myoneda@um.u-tokyo.ac.jp

The impact of climate change on human evolution is important and debating topic for many years. Since
2010, we have conducted a general joint project entitled “Replacement of Neanderthal by Modern Humans:
Testing Evolutional Models of Learning”, which based on a theoretical prediction that the cognitive ability
related to individual and social learning divide fates of ancient humans in the very unstable Late Pleistocene climate. This model predicts that the human populations which experienced a series of environmental
changes would have higher rate of individual learners, while detailed reconstructions of global climate change
have reported fluent and drastic change based on ice cores and stalagmites. However, we want to understandthe difference between anatomically Modern Human which survived and the other archaic extinct humans
including the European Neanderthals and Asian Denisovans.

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For this purpose the global synchronic change is not useful for understanding but the regional difference inthe amplitude and impact on biome of climate change is the information required. Hence, we invited a geophysicist using Global Climate Model (GCM) to reconstruct the climatic distribution and temporal change
in a continental scale. At the same time, some geochemist and geographers construct a database of local climate changes recorded in different proxies. At last, archaeologists and anthropologists tried to interpret the
emergence and disappearance of human species in Europe and Asia on the reconstructed past climate maps
using some tools, such as Eco-cultural niche model. Our project will show the regional difference in climatechange and related archaeological events and its impact on the evolution of learning ability of Modern Humans.

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Stadial and interstadial climates in the Late Pleistocene, as
simulated in experiments with the MIROC climate model
Wing-Le Chan1*, Ayako Abe-Ouchi1,2, Ryouta O’ishi3,1 andKunio Takahashi2

1. Atmosphere and Ocean Research Institute, The University of Tokyo, Japan; 2. Research Institute for Global Change, Japan Agency ofMarine-Earth Science and Technology, Japan; 3. National Institute of Polar Research, Japan. *Email: wlchan@aori.u-tokyo.ac.jp

The Late Pleistocene was a period which lasted from the Eemian interglacial period, approximately 126,000
years ago, to the start of the warm Holocene, 11,700 years ago, and was characterized mostly by widespreadglacial ice. Ice core data reveal abrupt climate changes, known as the Dansgaard-Oeschger events during
which relatively rapid warming in excess of 8℃ occurs over Greenland within the span of a few decades, followed by progressive cooling over 1,000 years or more. Some of the cool stadial states coincide with Heinrich events when icebergs broke off from the melting Laurentide ice sheet and ice rafted debris carried by thetraversing icebergs were deposited on the sea floors of the North Atlantic Ocean. The Late Pleistocene wasalso a period which saw Modern Humans spread throughout the world and other species of the same genus
such as the Neanderthals become extinct. Various hypotheses have been put forward to explain the extinction of Neanderthals, the last remains of which date back to about 30,000 years ago (Finlayson et al, 2006).
Among these is one which involves abrupt changes in past climate and the inability of Neanderthals to adapt
to such changes (Finlayson and Carrión 2006; Müller et al. 2011). The large discharge of freshwater into theNorth Atlantic Ocean during Heinrich events is thought to have greatly weakened the Atlantic meridional
overturning circulation, lowering North Atlantic sea surface temperatures (SST) with major environmental
consequences across parts of Europe such as cooling, aridification and a reduction in forest cover.

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In order to assess the effects of abrupt climate change on past human migration and on the extinction of certain species, it is first important to have a good understanding of the past climate itself. Previous investigations on abrupt climate changes during the last glacial period have used climate models forced under reduced
North Atlantic SST (e.g. Barron and Pollard 2002; Sepulchre et al. 2007). Some of these have also looked
into the vegetation response to these climate changes, either by using these reconstructed climate from atmosphere models or by using those from an atmosphere-ocean coupled model in which a freshwater flux is applied over the North Atlantic to simulate the reduction in SST there (Woillez et al. 2013).
In this study, we use three variants of ‘The Model for Interdisciplinary Research on Climate’, or known simply as MIROC (K-1 Model Developers 2001), a global climate model, for a time slice experiment within the
Late Pleistocene: two mid-resolution models (an atmosphere model and a coupled atmosphere-ocean model)
and a high-resolution atmosphere model. To obtain a fuller picture, we also look at a cool stadial state as
obtained from a ‘freshwater hosing’ coupled-model experiment, designed to mimic the effects of freshwater
discharge in the North Atlantic, as applied to a mid-glacial climate state. We use the sea surface temperature
response from this experiment to drive the atmosphere models. The surface air temperatures exhibit an antisymmetric pattern whereby cooling is seen across most of the northern hemisphere and weak warming is seenin parts of the southern hemisphere (Fig. 1). This is accompanied by drier climates in the north and wetterclimates in the south, in particular over the Southern Atlantic Ocean and northeast South America, indicative of a southward shift of the Intertropical Convergence Zone. More detailed features of the model-simulated climates are discussed and the results are compared with some available proxy data to elucidate wheremodel simulations show good agreement. The output from these climate model experiments are used todrive the Lund-Potsdam-Jena dynamic global vegetation model (Sitch et al. 2003) so that we can investigate

157

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changes in vegetation when the climate enters the cold stadial phase. In general, there is a reduction in forestover most of Europe, especially over the British Isles, north of central Europe and north of the Caucasus (Fig.
2). Much of this is replaced by grassland. On the other hand, there is also a small area stretching from theeastern coast of the Iberian Peninsula, across the islands in the western Mediterranean, to the western coastof Italy where forest cover increases.
References
Barron E, Pollard D (2002) High-resolution climate simulations of Oxygen Isotope Stage 3 in Europe. Quaternary Research 58:
296-309.
Finlayson C, Pacheco FG, Rodriguez-Vidal J, Fa DA, Gutierrez López JM, Santiago Pérez A, Finlayson G, Allué E, Baena
Preysler J, Cáceres I, Carrión SJ, Fernández Jalvo Y, Gleed-Owen CP, Jiménez Espejo FJ, López P, López Sáez JA, Riquelme
Cantal JA, Sánchez Marco A, Giles Guzman F, Brown K, Fuentes N, Valarino CA, Villalpando A, Stringer CB, Martínez Ruíz F,
Sakamoto T (2006) Late survival of Neanderthals at the southernmost extreme of Europe. Nature 443: 850-853.
Finlayson C, Carrión JC (2006) Rapid ecological turnover and its impact on Neanderthal and other human populations. Trends
in Ecology and Evolution 22: 213-222.
K-1 Model Developers (2004) K-1 Coupled GCM (MIROC) description, K-1 technical report (Hasumi H, Emori S eds), Center for Climate System Research, The University of Tokyo, Japan.
Müller UC, Pross J, Tzedakis PC, Gamble C, Kotthoff U, Schmiedl G, Wulf S, Christianis K (2011) The role of climate in the
spread of Modern Humans into Europe. Quaternary Science Reviews 30: 273-279.
Sepulchre P, Ramstein G, Kageyama M, Vanhaeren M, Krinner G, Sánchez-Goñi M-F, d’Errico F (2007) H4 abrupt event andlate Neanderthal presence in Iberia. Earth and Planetary Science Letters 258: 283-292.
Sitch S, Smith B, Prentice IC, Arneth A Bondeau A Cramer W, Kaplan JO, Levis S, Lucht W, Sykes MT, Thonicke K, Venevsky
S (2003) Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation
model. Global Change Biology 9: 161–185.
Woillez M-N, Kageyama M, Combourieu-Nebout N, Krinner G (2013) Simulating the vegetation response in western Europe
to abrupt climate changes under glacial background conditions. Biogeosciences 10: 1561-1582.

Fig. 1 The annual mean change in surface air temperature and precipitation when freshwater hosing conditions are applied to
an atmospheric model.

Fig. 2 Forest, grass and bare ground fractional changes across Europe, as calculated by a dynamical vegetation model.
Positive values correspond to increases when freshwater hosing conditions are applied.

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Geochemistry of Neanderthal and Modern Human archeologicalsites: Evidences of paleoclimatic variations and Homo s. activities
Francisco J. Jimenez-Espejo1*, Guadalupe Monge Gómez2,
Miguel Cortés Sanchez3, Clive Finlayson4,5, Joaquin Rodriguez-Vidal6 andAntonio García-Alix7

1. Depart. Biogeochemistry (JAMSTEC), Japan; 2. Dpto. Cristalografía, Mineralogía y Química Agrícola, Universidad de Sevilla, Spain; 3. Dpto.
Prehistoria y Arqueología, Facultad de Geografía e Historia, Universidad de Sevilla, Spain; 4. The Gibraltar Museum, UK; 5. Department ofSocial Sciences, University of Toronto, Canada; 6. Dpto. Geodinámica y Paleontología, Facultad de Ciencias Experimentales, Huelva, Spain;
7. School of Geographical and Earth Sciences, University of Glasgow, UK. *Email: fjjspejo@jamstec.go.jp

Introduction
On current evidence, southern Iberia is the region where the extinction of the last Neanderthal populations took
place (Finlayson et al. 2006). This information has been obtained from different archeological places located incave and rock shelters like those where is based our study: Gorham´s, Bajondillo, El Pirulejo and Vanguard. Thesekind of depositional environments contain major information about past Homo activities, paleoenviromental
sequences and faunal/floral evolution. Nevertheless, cave deposits are formed by a combination of erosional,
structural, karstic, biological and diagenetic process making these systems very complex from a geological point ofview: each cave presents its own unique features. Inorganic geochemistry can help us to resolve this complexity by
distinguishing the composition and evolution for the three major inputs that compose cave deposits: inorganic,organic and diagenetic.
Inorganic input to caves is associated with autochthonous minerals originating through calcite dissolution (theso-called ‘‘terra rossa’’), and external inputs. External inputs are mainly related to: (a) aeolian input in southern
Iberian coastal outcrops (Gorham´s, Vanguard and Bajondillo); and (b) surface or near-surface materials
occurring around the cave, usually after suffering a certain degree of pedogenesis, in those more continental (ElPirulejo). Organic deposits are associated with faunal/floral remains, guano, artifacts and combustions remains(hearths), among others. Diagenesis promotes new minerals formation (calcite, silica, etc) and also can affectelement distribution along soil profile, among other phenomena. These inputs during the conditions of formation
can be characteristic and can generate a distinctive geochemical imprint.

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This study aims to discover formation process, local disturbance, lateral progressive variations (e.g., entrance
to deep location and hearth distances), diagenetic effect, identify new proxies for anthropogenic activities andevaluate contamination levels associated to these activities.
At the continental site of El Pirulejo an Upper Paleolithic sequence has been described, including Solutrean
and Magdalenian (from 18 to 10 kyr) and Bronze Age activities. Preliminary results at El Pirulejo indicatethat Bronze Age archeological levels show high contents in Pb and Cu. These elements are almost absent in
Paleolithic levels and this presence between those is a clear indication of post-depositional alteration of theseolder levels associated to burials digging during the Bronze Age.
At coastal sites we can reconstruct the presence of Saharan aeolian input by the La/Lu ratio. The La/Lu ratio has
been specifically used to discriminate aeolian inputs of the African craton from those of the European margin(Cortés-Sánchez et al. 2012). The highest La/Lu values develop during Heinrich events that represent cold andarid episodes in this region which are associated with occupational hiatuses, affecting both, Neanderthals andModern Human populations ( Jimenez-Espejo et al. 2013). In these locations detrital proxies (K/Al and Mg/Al)

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show homogeneous values along similar stratigraphic levels, indicative of lower bio-turbation in the deep part ofthe cave, where a sub-horizontal stratigraphy has been preserved (Finlayson et al. 2006).
Samples obtained from clearly defined hearths have been compared with experimental fires allowing recognitionof a batch of elements associated to ash input. Very high values have been reached at specific points, allowing us
to indicate the earliest presence of anthropogenic mediated contamination in the geological record associated to
Neanderthals (Gorham´s Cave) although they can also be associated with diagenetic accumulations. In addition,
available data indicate certain similarity in rare earth element composition between ash elements and those
associated to guano and discrimination between these sources requires further work. In any case it is remarkablethat the ash/carbon related element exhibits a homogeneous dispersion along levels associated with hearths(Vanguard Cave), pointing to ash redistribution along the cave by Neanderthals, as observed in other locations(Vallverdú et al. 2010).
References
Cortés-Sánchez M, Morales-Muñiz A, Simón-Vallejo MD, Lozano-Francisco MC et al (2012) Earliest known use of marine
resources by Neanderthals. PLoS One 6: e24026.
Finlayson C, Giles Pacheco F, Rodríguez-Vidal J, Fa DA, Gutiérrez López JM et al (2006) Late survival of Neanderthals at thesouthernmost extreme of Europe. Nature 443: 850–853.
Jiménez Espejo FJ, Rodríguez Vidal J, Finlayson C, Martínez Ruiz F, Carrión JS, García Alix A et al (2013) Environmental conditions and geomorphologic changes during the Middle-Upper Paleolithic in the southern Iberian Peninsula. Geomorphology
180-181: 205-216.
Vallverdú J, Vaquero M, Cáceres I, Allué E, Rosell J, Saladié P, Chacón G, Ollé A, Canals A, Sala R, Courty MA, Carbonell E(2010) Sleeping activity area within the site structure of archaic human groups: Evidence from Abric Romaní Level N combustion activity areas. Current Anthropology 51: 137-145.

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Endocast surface reconstruction from CT images

Takashi Michikawa1*, Masaki Moriguchi2, Hiromasa Suzuki3,
Naomichi Ogihara4, Osamu Kondo3 and Yasushi Kobayashi5

1. Osaka University, Japan; 2. Chuo University, Japan; 3. The University of Tokyo, Japan; 4. Keio University, Japan; 5. National Defense
Medical College, Japan. *Email: michikawa@acm.org

Introduction
An endocast is the internal cast of a cranium where a brain shape is stored. Extracting geometric models of
endocasts is an important issue in anthropology, because endocasts give us various useful anthropologicalinformation such as brain shapes and their volumes. We usually use CT scanners, non-destructive measurement
devices in order to extract geometric data of crania. The scanned result or CT images contain scalar value (CTvalue) corresponding to the materials. Since CT value is proportional to material density, the surfaces can beextracted by common isosurface extraction methods (Lorensen 1987) with an isovalue. However, this approachis difficult to extract endocast surfaces, because the isosurfaces contains whole surface of crania and an additional
manual operation is required for extracting only endocast surfaces.
We introduce a computational method for extracting endocast shapes from CT images of crania. From ourobservation, we found the endocast is the largest empty space in a cranium. We use this heuristics for automatic
endocast surface extraction problem. Our method is based on the watershed-based segmentation (Vincent 1991)
using distance fields. We first compute distance fields from a binary images of a cranium. Next, we find initial
seed voxels using the distance field so that a part of endocast region and corner voxels are individually selected.
Finally, the watershed segmentation is applied to the binary images and they are classified into crania part,
endocast part and background part. We use this segmentation result for polygonization of the endocast. We also
demonstrate our method can extract polygonal meshes from CT images of various kinds of crania including,monkey and assembled fossil crania.

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Algorithm
Our endocast extraction method consists of four steps (Fig. 1) and their details are described below.
The first step is the determination of crania objects from CT images. This consists of two steps; binarizationand noise removal. Binarization classifies the input images into a crania shape and background and we binarize
this with an isovalue t. The results often contain noise due to scanning artifacts and we apply labelling to findconnected components to the results and the largest component is extracted as foreground voxels or the cranium
object.
The next step is to extract an endocast voxels from the binary images by using watershed algorithm using distancefields (Fig. 2). We first compute distance field (Lohmann 1998) from the binary image (Fig. 2 (b)). Next, we findinitial seed box v such that d (v) > k max d (v) where d (v) and k denote the distance value at v and user-given
parameter respectively. We choose k so that a few endocast voxels and background voxels are selected. In addition,
we also select corner voxels of the binary images as initial seed voxels because boundary voxels must become
single component. We apply the labeling to the initial seed voxels and the two components are labelled (Fig. 2(c)).
Then, the watershed algorithm is applied to expand initial seeds using distance value. When the algorithm is161

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terminated, the input volume data is decomposed into cranium part (white), endocast part (red) and backgroundpart (yellow) shown in Fig. 2(d).
The final step is polygonization of endocast region classified in the previous step. We here apply Marching cubes
to masked cells so that any unmasked cells are not polygonized. The isovalue is same as that used in the firstbinarization step.
Results and discussion
Experimental results are shown in Fig. 3. In each example, an isosurface of CT images and the result are drawn
in gray and red respectively. We confirmed that the endocast surfaces could be extracted from various typesof examples, including a monkey (Fig. 3 (b)) and a fossil cranium (Fig. 3 (c)). Note that sub-voxel accuracy isguaranteed since polygonization is based on Marching cubes.
The statistics of the experiments are summarized in Table 1. All the experiments are tested on Windows PC(Intel Core i7-3930K CPU + 64 GB RAM). Each computation is automatic and computation time is about ten
minutes. Our method has two major parameters: isovalue t and scaling factor k. In particular, we have to select s
carefully so that initial seed voxels are decomposed into two components. For instance, base of the skull is often
lost in fossil cranium models and the holes are relatively large. In such cases, we need to select larger scaling
parameter. This parameter tuning is easy by using rendering results.
Our method has two limitations. The first limitation is that surface quality depends on the quality of binarization.
Since the current implementation uses a single threshold, thin features are often lost by binarization. The second
limitation is that our method is difficult to control surface boundary. Our method is designed for extracting theinner surface of a cranium and unnecessary region is extracted from incomplete models such as fossil crania (Fig.
3(c)), additional removing operation may be required to extract endocast surfaces. However, the task becomesmuch simpler than conventional approaches.
Conclusion
We have introduced a method for computing endocast polygons from CT images of crania. Our method extracts
the endocast region by watershed algorithm. Polygonization of endocast surfaces is computed by Marching cubes
algorithm with the segmentation result as mask images. Experimental results showed that our method can extract
endocast polygons from various kinds of crania with few parameter settings. As a result, the task of extracting
endocast becomes much simpler.
There are some future issues for this research. Main issue is to introduce non-uniform polygonization fromvolume data. Particularly, per-voxel threshold is required for high-quality binarization. The other issue is to findinitial seed automatically.
References
Lorensen WE, Cline HE (1987) Marching cubes: A high-resolution 3d construction algorithm. ACM Computer Graphics 21 (4):163-169.
Vincent L, Soille P (1991) Watersheds in digital spaces: An efficient algorithm based on immersion simulations. IEEE Transactions on Pattern Analysis and Machine Intelligence 13(6): 583-598.
Lohmann G (1998) Volumetric Image Analysis. Wiley.

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Fig. 1 An overview of our method. (a) Input volume data (512x512x301 voxels). (b) Binarization result. (c) Segmentation result(red). (d) Endocast polygon of (a) (803,296 faces).

Fig. 2 Endocast segmentation based on watershed algorithm. (a) Binary image. (b) Distance field. (c) Initial seed voxels (red/yellow) computed from distance value. (d) Segmentation result.

Fig. 3 Results.

Table 1 Statistics for the experiments.

Data
Name
KUMA (Fig. 1)
K2589 (Fig. 3 (a))
M15 (Fig. 3 (b))
Mladec1 (Fig. 3 (c))

ParametersSize
512x512x301
512x512x399
512x512x253
512x512x360

t
25
45
230
910

k
0.35
0.85
0.15
0.35

#faces803,296
1,052,296
182,280
1,137,492

Time [s]
304.5
443.7
585.7
377.7

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Reconstruction of Qafzeh 9 braincase and its endocast
morphology
Osamu Kondo1*, Daisuke Kubo2, Hiromasa Suzuki3 and Naomichi Ogihara4
1. Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Japan; 2. School of Health and Physical
Education, University of Tsukuba, Japan; 3. Department of Precision Engineering, The University of Tokyo, Japan; 4. Department of
Mechanical Engineering, Faculty of Science and Technology, Keio University, Japan. *Email: kondo-o@biol.s.u-tokyo.ac.jp

Introduction
Paleoneurology is an important field in the study of human evolution as it contributes important information
to other areas of paleoanthropology. Variations in the size and shape of endocasts help in differentiating among
fossil species. Moreover, comparisons of the size and shape of the fossil endocasts provide considerable linksof the hard evidence to behavioral and cognitive differences among fossil species, which might have drove thespeciation between the taxa. In this context, C01 team of the project RNMH (Replacement of Neanderthals byModern Humans) aims to present reasonable ways for reconstruction of the fossil hominids in order to estimate
the endocasts and the brains, and also aims to make possible inferences for behavioral and cognitive differences
between Neanderthals and Modern Humans, based on the estimated endocasts and brains.

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Preliminary results and the scope
In this paper, we describe the endocast of Qafzeh 9 and compare the size and shape of it with those of Modern Humans and several fossil hominids. The endocast reconstruction of the Qafzeh 9 has been accomplishedby ourselves (Kondo et al. 2014), where a semi-virtual reconstruction was performed by using sequential CT
images of the present cranial reconstruction with their planar resolution of 0.41 mm/pixel and the slice thickness of 0.5 mm. The process followed that of Kubo and Kono (2011), i.e. after extracting the cranial modelfrom the CT images by using a threshold between the bone and air and making a physical mold of the endocranium with 3D printer, we filled the gap and repaired the lacking portions of the inner cranial surface withclay. The more complete surface model of the endocast was produced from re-scanning images of the repaired
mold. At the same time, with assessment of the degree of R-L asymmetry of the original fossil cranium,
we have tried to correct the observed degree of deformation or distortion along the most variant direction
against the normal R-L asymmetry of Modern Human crania. The assessment suggested that the present reconstruction of the Qafzeh 9 cranium, possessing a significant degree of R-L asymmetry, might have sufferedfrom distortion (Kondo et al. 2014).
Directly measured endocranial volumes, 1411 cc for the original reconstruction and 1477 cc for the morphed
version, respectively (Fig. 1) were smaller than previously published data (1531 cc, Holloway et al. 2004) using least-square regression equations. After morphing, the endocast was higher at the middle cranial fossa
and narrower at the frontal lobe, both characteristics falling within the range of recent human variation.
Although the reconstructed Qafzeh 9 endocasts of both the original and morphed versions fit into the endocranial evolution among lineages of the genus Homo, many fossil crania may have undergone a substantial
degree of taphonomic deformation. Therefore, application of possible corrections or hypothesis of plausible
taphonomic scenarios should be part of a reasonable assessment of a single precious fossil specimen.

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Qafzeh 9 was discovered and excavated by B. Vandermeersch in 1969 from the Qafzeh Cave in Israel. It was
associated with Middle Paleolithic assemblage of the Levalloiso-Mousterian culture (Vandermeersch 1981).

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In spite of the older chronological age over 90,000 years and the older type of associated lithic assemblage(Tabun C type in Middle Paleolithic), the human skeletal morphology from the Qafzeh Cave is quite modern and contrasted to those of contemporary or even younger classic Neanderthals. According to his firstdescription, the Qafzeh 9 individual is an adult female, whose cranial bones were crashed and heavily fragmented, but the whole cranium was well restored and reconstructed (Vandermeersch 1981). The endocranialfeature was only described on the parietal meningeal vessels because no endocasts have been made for thisspecimen. Therefore, even now we know the present reconstruction exhibits possible distortions both in theneurocranium and the face, it is worth describing the original version of the endocast and then comparing thesize and shape of both the original and the morphing version with those of modern and fossil humans.
References
Holloway RL, Broadfield DC, Yuan MS (2004) The human fossil record, vol. 3. Brain endocasts -The paleoneurological evidence.
Wiley-Liss.
Kondo O, Kubo D, Suzuki H, Ogihara N (2014) Virtual endocast of Qafzeh 9, a preliminary assessment of right-left asymmetry.
In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 2:
Cognitive and physical perspectives. Springer, New York, pp 183-190.
Kubo D, Kono RT (2011) Endocranial restoration and volume estimation of the Minatogawa IV cranium using micro-CT and
3D printer systems. Anthropological Science 119: 203-209.
Vandermeersch B (1981) Les hommes fossiles de Qafzeh (Israel). Editions du CNRS, Paris.

Fig. 1 Reconstruction of Qafzeh 9 endocast. org: original
version in the left, mor: morphing version (reduced R-Lasymmetry) in the right (modified from Kondo et al. 2014).

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Surface deformation of crania by optimizing rigidity

Masaki Moriguchi1*, Hiromasa Suzuki2, Takashi Michikawa3,
Naomichi Ogihara4 and Osamu Kondo5

1. Department of Information and System Engineering, Chuo University, Japan; 2. Department of Precision Engineering, The University ofTokyo, Japan; 3. Division of Sustainable Energy and Environmental Engineering, Osaka University, Japan; 4. Department of MechanicalEngineering, Keio University, Japan; 5. Department of Biological Science, The University of Tokyo, Japan. *Email: moriguchi@ise.chuo-u.ac.jp

Introduction
Shape deformation is a useful tool in geometric morphometrics, since it can be used to reconstruct missing parts
and transfer semi-landmarks. Thin-plate spline (TPS) warping (Bookstein 1989), which minimizes the bending
energy of transformations, is a predominantly used deformation method in geometric morphometrics. Missing
parts are reconstructed by transforming a reference model (that has no missing parts) to a model with missing
parts (Gunz et al. 2009). The warping function of TPS can be computed by solving dense linear systems of size (k+ 4) × (k + 4), where k is the number of interpolation points (i.e. landmarks).

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TPS warping can also be used to transfer semi-landmarks. Suppose we have two models A and B, where only
model A has semi-landmarks, and we want to transfer the semi-landmarks from A to B. The sliding semilandmark method (Bookstein 1997) positions the semi-landmarks on B so that the induced TPS warping has theminimum bending energy. The variables in this minimization problem are the coordinates of the semi-landmarks
on B and, since the semi-landmarks are needed to lie on B, this is a constrained minimization problem. Local
minima of the problem are computed by alternating between relaxed optimization and projection. In the relaxed
optimization step, the constraints that semi-landmarks lie on B are removed and, instead, each semi-landmark s
on B is allowed to move on the tangent plane of B at s. With these relaxed constraints, the optimization can now
be solved by solving linear systems. In the projection step, semi-landmarks that move away from B in the relaxed
optimization step are projected back to their closest points on B.
There are mainly two types of methods for shape deformation: space deformation and surface-based deformation.
Space deformation methods deform a surface by deforming a space that encloses the surface; the surface isimplicitly deformed according to the deformation of the space. TPS warping is a space deformation methodthat deforms the entire 3D space. Space deformation methods are robust in the sense that they can operate on
imperfect data such as point clouds, triangle soups and surface meshes with bad shape quality. Surface-based
deformation methods directly deform a surface. Variational surface-based deformation methods compute thedeformation by minimizing some deformation energy (such as bending energy) and they achieve high quality
shape-preserving deformation.

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Several linear variational methods that minimize linearized deformation were proposed in recent years (Botschand Sorkine 2008). They are fast, since the deformation is computed by solving (sparse) linear systems. However,they might cause artifacts in the deformed surface. As-rigid-as-possible (ARAP) surface deformation (Sorkine
and Alexa 2007) is a popular surface-based deformation method in geometric modeling that avoids the artifactsof linear variational methods. The ARAP surface deformation method computes a high quality shape-preserving
deformation by minimizing the ARAP deformation energy. Though its energy formulation is non-linear, it
is conceptually simple and easy to implement. In this work, we apply ARAP surface deformation to Modern
Human crania and compare the results with the ones obtained by TPS warping.

Poster Presentations

MethodAs-rigid-as-possible (ARAP) surface deformation finds a transformation that is locally a rigid transformation. The ARAP energy formulation is derived as follows (Sorkine and Alexa 2007). Suppose we have a surface mesh S whose vertex set is V and edge set is E. We denote the deformed surface mesh as S'. The ARAP
energy measures the local deviations from rigid transformations and is defined aswhere pi is the position of the i-th vertex of S, pi' is the position of the i-th vertex of S' and Ni is the set ofneighbor vertices of vertex pi. wi is the weight of vertex pi and wij is the weight of the edge connecting pi and
pj. Ri is the optimal local rotation that transforms the neighbors of pi to those of pi'. In shape deformation,
the original surface mesh S is fixed and some vertices under positional constraints in S' are also fixed. Thus,
the ARAP energy becomes the function of the coordinates of unconstrained vertices in S'. The ARAP deformation method computes these coordinates by minimizing the ARAP energy. Sorkine and Alexa (2007)
solve this minimization problem by a twostep alternating optimization:
・Fix the local rotations Ri (i ∈ V) and optimize the positions of unconstrained vertices in S'.
・Fix the positions of unconstrained vertices in S' and optimize the local rotations Ri (i ∈ V).
This optimization deals with Ri as if they are independent variables from pi and its neighbor vertices. Suppose we have a reference mesh A (that has no missing parts) and a mesh B with missing parts, where both A
and B have landmarks. To reconstruct missing parts, we compute a deformed reference mesh A' using ARAP
deformation under the following positional constraints: the position of each landmark on A' is set to thatof the corresponding landmark on B. By optimizing the positions of unconstrained vertices in A' using the
ARAP energy, we obtain a desired deformed mesh A'.
References
Bookstein FL (1989) Principal warps: Thin-plate splines and the decomposition of deformations. IEEE Transactions on PatternAnalysis and Machine Intelligence 11(6): 567-585.
Bookstein FL (1997) Landmark methods for forms without landmarks: Morphometrics of group differences in outline shape.
Medical Image Analysis 1(3): 225-243.
Botsch M, Sorkine O (2008) On linear variational surface deformation methods. IEEE Transactions on Visualization and Computer Graphics 14(1): 213-230.
Gunz P, Mitteroecker P, Neubauer S, Weber GW, Bookstein FL (2009) Principles for the virtual reconstruction of hominin crania. Journal of Human Evolution 57(1): 48-62.
Sorkine O, Alexa M (2007) As-rigid-as-possible surface modeling. In: Proceedings of the symposium on geometry processing.
pp. 109-116.

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Neural underpinning of creative thinking: A meta-analysis
for neuroimaging data
Naoki Miura1*, Yukako Sasaki2, Kunihiro Hasegawa3 and Hiroki C. Tanabe3

1. Faculty of Engineering, Tohoku Institute of Technology, Japan; 2. Institute of Development, Aging and Cancer, Tohoku University, Japan; 3.
Graduate School of Environmental Studies, Nagoya University, Japan. *Email: miura.naoki@tohtech.ac.jp

Introduction
The target of our research project is to investigate whether the cause of phenomenon for replacement of
Neanderthals by Modern Humans can explain according to difference of cognitive mechanism for learning
behavior from a viewpoint of neuroscience. For this purpose, we compare morphological differences of brain
between Neanderthals and Modern Humans by using computational data of fossil crania. And, differenceof neural representation for learning behavior could be discussed based on those morphological differences.
To discuss the difference of neural representation, it is necessary to depict a cognitive mechanism in whicha difference is predicted among Neanderthals and Modern Humans. And, it is also necessary to discuss thedifference of functional representation of that cognitive mechanism on brain region, that is obtained fromcontemporary human subjects, by projecting virtual Neanderthal brain.

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Cognitive mechanism of creative thinking is one of the convincing candidates which represent the differenceof Neanderthals and Modern Humans. It is considered that the drastic climate change repeatedly arose in
the age of replacement. In order to survive under the environment, creative and adaptive behaviour might berequired for various circumstances. Therefore, we expected that difference of cognitive mechanism to demonstrate
the creativity might be represented the difference of behaviour required for survival. And, the replacement
arose because the Modern Human who possessed capacity for creative and adaptive behaviour to survives inenvironmental perturbation. Recent reviews of neuroimaging for creativity (Dietrich and Kanso 2010; Arden etal. 2010) suggested that neural underpinning of creative thinking could not be explained depending on the single
cognitive process or cortical region. Thus, it is necessary to clarify a common neural underpinning of creativity
based on various kinds of experimental task to depict the creative behaviour.
The purpose of the present study is to depict the neural representation of creative behaviour to discuss thedifference of cognitive capability between Neanderthals and Modern Humans. For this purpose, we use a metaanalysis method for neuroimaging data to create functional map of creative behaviour. The research articles offunctional magnetic resonance imaging (fMRI) study for creative thinking are collected, and common corticalregions representing creative thinking is depicted by using the activation likelihood estimation (ALE) method
based on the reported cortical activation.
MethodIn the present study, we collected forty research articles describing neural basis of creative thinking measured
by fMRI. For details, twenty-six articles for fMRI investigations using verbal creative task, six articles for
fMRI investigations of creativity task using figure information, and eight articles for fMRI investigations using artistic creativity were collected.

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Based on activation tables which were reported in the collected articles, the three-dimensional coordinateof each activation peak was extracted. Montreal Neurological Institute (MNI) coordinate was used in whole

Poster Presentations

analysis. As the experimental task which produces creative activity, various kinds of experimental task wereused in each fMRI measurement. Therefore, We extracted activation peaks of subtraction analysis between
experimental condition with creative thinking and either conditions of lower control condition or resting
condition, in order to describe the cortical activity common to various creative activities. The coordinate ofactivation peak which was reported using Talairach space was converted to MNI space using the MTT transform (Lancaster et al. 2007). Consequently, 473 activation peaks were extracted.
A meta-analysis for activation peaks was performed using GingerALE software (BrainMap.org; http://www.
brainmap.org/ale/) which provide common activation maps from dataset using Activation Likelihood Estimation (ALE) method (Eickhoff et al. 2009, 2012; Turkeltaub et al. 2012). In order to estimate the common
activation pattern of creative behavior, ALE method was performed considering the extracted activation peals
as one dataset. The less conservative mask was used because it was for taking into consideration fluctuationof the activation peak located on the surface region of cerebrum. As a result of masking process, thirteen activation peaks were discarded. The ALE map and the P value on each voxel were calculated, and statisticalthreshold for ALE map was set by p < 0.01 using False-Discovery Rate (Laird et al. 2005).
Result and discussion
As a result of meta-analysis, cortical regions with statistically significant ALE value were observed in manydifferent cortical regions (Fig. 1). As the significant clusters in the frontal region, eight clusters were observedon left lateral prefrontal region, two clusters were observed on right lateral prefrontal region, and two clusters
were observed on medial prefrontal region. In another region, two clusters were observed on left parietal region, four clusters were observed on left temporal region, four clusters were observed on occipital region, andtwo clusters were observed on right cerebellum.
As an overall trend, the activation clusters were observed by predominance in the left hemisphere. This trend
was remarkable also in the prefrontal area. The left prefrontal area is strongly related to linguistic processing.
Since many of experimental tasks which were used in collected articles were used creativity test on verbal
information, the activation cluster in left prefrontal cortex was reflecting verbal processing. But, it was also
observed that cortical activations during artistic creativity tasks were involved in the ALE value of this cluster, the left prefrontal cortex is synthetically important for creative thinking. That result is consistent with thesuggestion in previous reviews of creativity (Dietrich and Kanso 2010; Arden et al. 2010).
Significant clusters were also observed in the left parietal and temporal cortex and right cerebellar posterior
lobule. In our group’s research about a reconstruction of the virtual Neanderthal brain using computational
anatomy (Tanabe 2014), it was presumed that these regions had morphological differences between Neanderthals and Modern Humans. That is, the volume of the parietal cortex and cerebellum of virtual Neanderthal
brain might be smaller than that of Modern Human, on the other hand the region from a posterior temporalto occipital cortex might be larger in virtual Neanderthal brain. When the morphological and functional
differences are taken into consideration together, the difference of creative thinking between Neanderthals
and Modern Humans may be inferred from the morphological difference on cognitive network entire brain
representing creativity. Thus, the present result provides a possible data to discuss the difference of cognitive
mechanism for learning behavior.
References
Arden R, Chavez RS, Grazioplene R, Jung RE (2010) Neuroimaging creativity: A psychometric view. Behavioural Brain Research 214(2): 143-156.
Dietrich A, Kanso R (2010) A review of EEG, ERP, and neuroimaging studies of creativity and insight. Psychological Bulletin
136(5): 822-848.

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Eickhoff SB, Laird AR, Grefkes C, Wang LE, Zilles K, Fox PT (2009) Coordinate-based activation likelihood estimation metaanalysis of neuroimaging data: A random-effects approach based on empirical estimates of spatial uncertainty. Human Brain
Mapping 30(9): 2907-2926.
Eickhoff SB, Bzdok D, Laird AR, Kurth F, Fox PT (2012) Activation Likelihood Estimation meta-analysis revisited. Neuroimage 59(3): 2349-2361.
Laird AR, Fox PM, Price CJ, Glahn DC, Uecker AM, Lancaster JL, Turkeltaub PE, Kochunov P, Fox PT (2005) ALE metaanalysis: Controlling the false discovery rate and performing statistical contrasts. Human Brain Mapping 25(1): 155-164.
Lancaster JL, Tordesillas-Gutiérrez D, Martinez M, Salinas F, Evans A, Zilles K, Mazziotta JC, Fox PT (2007) Bias between
MNI and Talairach coordinates analyzed using the ICBM-152 brain template. Human Brain Mapping 28(11): 1194-1205.
Tanabe HC (2014) Difference of brain function between Neanderthals and Modern Humans based on the cranial morphometry.
Proceeding of the 9th Conference on RNMH project 15.
Turkeltaub PE, Eickhoff SB, Laird AR, Fox M, Wiener M, Fox P (2012) Minimizing within-experiment and within-group effects in Activation Likelihood Estimation meta-analyses. Human Brain Mapping 33(1): 1-13.

Fig. 1 Significant activation clusters which are commonly observed during creative thinking tasks.
Z-value indicates the value of MNI coordinate on the z-axis from the origin towards the top of the
head.

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fMRI Experiment about executive function by difference oflearning methods: Using Tower of Hanoi
Yukinobu Hoshino1*, Keita Mitani2, Naoki Miura3, Hiroki C. Tanabe4 andKenji Nagai5

1. School of Systems Engineering, Kochi University of Technology, Japan; 2. Kochi University of Technology, Japan; 3. Faculty of Engineering,Tohoku Institute of Technology, Japan; 4. Graduate School of Environmental Studies, Nagoya University, Japan; 5. Tohoku University of Art
and Design, Japan. *Email: hoshino.yukinobu@kochi-tech.ac.jp

Introduction
In the previous report, we measured and analyzed 3D motion data by subject that has expert skills of therecurrent Levallois method. Execution of stone-tool production was shown to concern with “social learning”
and “individual learning” by our results. In this study, our purpose is to verify effect of executive function bysocial learning or individual learning from fMRI analysis. Executive function is defined as “the ability to retain
a relevant stance for the goal of future”, and consists of (1) goal formulation, (2) planning, (3) carrying out goaldirected plans and (4) effective performance (Muriel 1982). Then, executive function is lower brain functions thatconsist division attention, management of tasks, conversion ability to think and inductive guess. Execution ofstone-tool production in MRI is impossible, so we select the Tower of Hanoi as the task for executive function
analysis in this experiment (Humes et al. 1997). In this paper, we explain the experiment environment and theresults of fMRI analysis and task performance data.
Experiment

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This experiment was used our designed Tower of Hanoi task in fMRI. Purpose of this experiment is to verify
the effect of learning method using fMRI analysis. Subjects were three men in their twenties. In order toanalyze the effect of learning methods, three men played either of three groups of “non learning”, “individual
learning” and “social learning”. Non learning group did not have learning period in the experiment. Individual learning group had learning period that is practice of Tower of Hanoi. Social learning group had learning
period with teacher. Fig. 1 shows this experiment time schedule of each group.
Subjects performed the Tower of Hanoi task of onscreen using controller in MRI. This task has three to six
disk of Tower of Hanoi. This experiment has two sessions (Session:1, Session:2). A session is consisted ofeight Task parts (Task1 & Task2 * three to 6 disks), eight Repeat parts (Task1 & Task2 * three to 6 disks) andsome rest parts. Fig. 2 shows detail time sequence of a session.
Task and Repeat have two parts of Tower of Hanoi task by three to six disks. In Task parts, Subjects performed a repeat in part time (40 second). In Repeat parts, the screen was displayed his moved disk of Task
part (40 second). Therefore, subjects looked the screen, and performed same control. In rest parts, subjects did
not perform or think (15 second). In this fMRI experiment, we measurement the average performance time
and task completed count. In the next heading, we report some results about fMRI analysis, performancemeasurement.
Results
Average performance times of all subjects are shown in Fig. 3.

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About tendency of non learning subject results, average performance times are almost constant in all tasks.
Individual learning subject and social learning subject results of average performance time were shown eachtendency. About tendency of individual learning subject, average performance times of Session:2 were shorter
than Session:1. In particular, tasks of six disks have notable difference. Average performance times of social
learning subject also were short at Session:2. However, times of Session:2 six disks were longer than Session:1. We think that this reason is delay by recalled of learned subject. From these results, we think that non
learning subject did not learn about task in experiment, but it is evident because he did not have learning
period. Performance of individual learning subject and social learning subject improve after the each learning,
so we think that individual learning subject and social learning subject progress learning.
Next, fMRI analysis data of individual learning subject and social learning subject are shown in Fig. 4.
Results of Fig. 4 are difference of Session:1 and Session:2 at six disks. Learning progresses of individual
learning subject and social learning subject were observed from results of performance measurement, but differences by effect of learning are not observed from fMRI data results. We think that this cause has someproblem to task design. One problem is that we need observe until end of learning. Then, we need conduct
experiment with more subjects. Other detail results will report at this conference.
References
Lezak MD (1982) The problem of assessing executive functions. International Journal of Psychology 17: 281-297.
Humes GE, Welsh MC, Retzlaff P, Cookson N (1997) Towers of Hanoi and London: Reliability of two executive function tasks.
Assessment 4(3): 249-257.

Fig. 1 Experiment time schedule of each group.

Fig. 3 Average performance times of each task.

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Fig. 2 Detail time sequence of a session.

Fig. 4 fMRI analysis data of individual learning subject (a), social learning subject (b).

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Brain structural and functional analyses for motivation tolearn, fatigue and cognitive function
Kei Mizuno1-3

1. Pathophysiological and Health Science Team, RIKEN Center for Life Science Technologies, Japan; 2. Department of Medical Science on
Fatigue, Osaka City University Graduate School of Medicine, Japan; 3. Osaka City University Center for Health Science Innovation, Japan.
Email: keimizuno@riken.jp

Differences in learning abilities between ancient and modern humans can be demonstrated by using functional and structural neuroimaging methods to identify the brain regions that are involved in learning inmodern humans, and then by using fossil reconstruction to compare these regions between modern and ancient humans. The research term C02 (Functional Mapping of Learning Activities in Archaic and Modern
Human Brains), part of the study project for Replacement of Neanderthals by Modern Humans (RNMH),
has focused on factors for enforced learning such as creativity and social cognition.

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The motivation to learn is one such factor. We have used functional magnetic resonance imaging (fMRI) in
order to investigate the neural substrates of the motivation to learn in modern humans. We confirmed thatsense of accomplishment was an intrinsic reward and was associated with the motivation to learn (Yoshida etal. 2008). The putamen, including the striatum, was strongly activated by sense of accomplishment, and putamen activation was positively associated with a motivation to learn score derived from a questionnaire (Mizuno
et al. 2008). Academic rewards for motivational arousal include a sense of accomplishment and competence,
self-efficacy, and acceptance by others (Deci et al. 1991, 1996). Perception of a good reputation by others is
processed by the medial prefrontal cortex and the striatum (Izuma et al. 2008). These results suggest that thestriatum plays a crucial role in the motivation to learn. Structural analysis of the brain in modern humans
revealed that the motivation to learn score was positively correlated with the volume of the right putamen(Takeguchi et al. 2014), indicating that both the volume and the neural activity of the striatum are closely associated with the motivation to learn.
There is a negative correlation between the motivation to learn and fatigue, indicating that these phenomena
are two sides of the same coin, and that accumulation of fatigue negatively reinforces not only motivation tolearn, but also learning. The medial orbitofrontal cortex was associated with fatigue sensation using positronemission tomography (Tajima et al. 2010), and the activity of the medial orbitofrontal cortex, assessed using
fMRI, increased with time spent on a fatigue-inducing task. In addition, activation of the medial orbitofrontal cortex was attenuated by the perception of an intrinsic reward, suggesting that intrinsic reward modulates
the extent of fatigue alleviation.

174

To clarify the neural bases of motivation to learn and fatigue, we have determined what cognitive functions
are correlated with the level of motivation to learn and/or the level of fatigue. When interpreted in light ofthe differences in brain morphology and function between modern and ancient humans, the neural substratesof these cognitive functions may provide information on the apparent differences in learning ability. In elementary and junior high school students, motivation to learn was associated with divided-attention performance, evaluated using the kana pick-out test (Mizuno et al. 2011a). Fatigue, which was evaluated using task
E on the modified advanced trail making test (Mizuno et al. 2011b), was negatively associated with performance on the divided-attention task and with switching attention. Children and adolescents with chronic fatigue syndrome (CFS), which is characterized by profound and disabling fatigue for at least 6 months (Fukuda
et al. 1994), showed poor performance on the divided-attention and switching-attention tasks (Tomoda et al.
2007; Kawatani et al. 2012). The ability to divide and switch attention dramatically develops from elementary

Poster Presentations

to junior high school (Mizuno et al. 2011c). These results suggest that motivation to learn and fatigue influence the age-dependent development of brain function. Using fMRI, we revealed that in young adults, thebrain regions involved in the kana pick-out test are the left dorsal inferior frontal gyrus and the superior parietal lobule (Mizuno et al. 2012). Another fMRI study showed that the dorsolateral prefrontal cortex is crucial for the process of switching attention in the trail making test (Moll et al. 2002). A study performed using
voxel-based morphometry demonstrated volume reduction of the bilateral dorsolateral prefrontal cortices inadult patients with CFS, and the reduction in volume in the right prefrontal cortex was associated with theseverity of fatigue (Okada et al. 2004). Our recent study showed that the volume of the bilateral dorsolateralprefrontal cortices was also associated with fatigue in healthy adults.
These findings highlight the regions of the brain, including the striatum and medial orbitofrontal cortex, themedial prefrontal cortex, the inferior frontal gyrus, the superior parietal lobule, and the dorsolateral prefrontal
cortex, that are of interest in relation to the motivation to learn, fatigue, and cognitive functions affected bymotivation and fatigue in modern humans. Morphological and functional comparisons of these regions between modern and ancient humans are expected to help elucidate differences in learning abilities. We expect
these outcomes to contribute to the achievements of the RNMH project.
References
Deci EL, Vallerand RJ, Pelletier LG, Ryan RM (1991) Motivation in education: The self-determination perspective. Educational
Psychologist 26: 325-346.
Deci EL, Ryan RM, Williams GC (1996) Need satisfaction and the self-regulation of learning. Learning and Individual Differences 8: 165-183.
Izuma K, Saito DN, Sadato N (2008) Processing of social and monetary rewards in the human striatum. Neuron 58: 284-294.
Fukuda K, Straus SE, Hickie I, Sharpe MC, Dobbins JG, Komaroff A (1994) The chronic fatigue syndrome: A comprehensive
approach to its definition and study. International Chronic Fatigue Syndrome Study Group. Annals of Internal Medicine 121:
953-959.
Kaneko M (1996) A neuropsychological index of aging data from 9000 elderly people. Nippon Ronen Igakkai Zasshi 33: 811815. (In Japanese)
Kawatani J, Mizuno K, Shiraishi S, Takao M, Joudoi T, Fukuda S, Watanabe Y, Tomoda A (2011) Cognitive dysfunction andmental fatigue in childhood chronic fatigue syndrome--A 6-month follow-up study. Brain and Development 33: 832-841.
Mizuno K, Tanaka M, Ishii A, Tanabe HC, Onoe H, Sadato N, Watanabe Y (2008) The neural basis of academic achievement
motivation. Neuroimage 42: 369-378.
Mizuno K, Tanaka M, Fukuda S, Imai-Matsumura K, Watanabe Y (2011a) Relationship between cognitive function and prevalence of decrease in intrinsic academic motivation in adolescents. Behavioral Brain Functions 7: 4.
Mizuno K, Tanaka M, Fukuda S, Imai-Matsumura K, Watanabe Y (2011b) Relationship between cognitive functions and prevalence of fatigue in elementary and junior high school students. Brain and Development 33: 470-479.
Mizuno K, Tanaka M, Fukuda S, Sasabe T, Imai-Matsumura K, Watanabe Y (2011c) Changes in cognitive functions of students
in the transitional period from elementary school to junior high school. Brain and Development 33: 412-420.
Mizuno K, Tanaka M, Tanabe HC, Sadato N, Watanabe Y (2012) The neural substrates associated with attentional resources anddifficulty of concurrent processing of the two verbal tasks. Neuropsychologia 50: 1998-2009.
Moll J, de Oliveira-Souza R, Moll FT, Bramati IE, Andreiuolo PA (2002) The cerebral correlates of set-shifting: An fMRI studyof the trail making test. Arq Neuropsiquiatr 60: 900-905.
Okada T, Tanaka M, Kuratsune H, Watanabe Y, Sadato N (2004) Mechanisms underlying fatigue: A voxel-based morphometricstudy of chronic fatigue syndrome. BMC Neurology 4: 14.
Tajima S, Yamamoto S, Tanaka M, Kataoka Y, Iwase M, Yoshikawa E, Okada H, Onoe H, Tsukada H, Kuratsune H, Ouchi Y,
Watanabe Y (2010) Medial orbitofrontal cortex is associated with fatigue sensation. Neurology Research International 2010:
671421.
Takeuchi H, Taki Y, Nouchi R, Sekiguchi A, Kotozaki Y, Miyauchi CM, Yokoyama R, Iizuka K, Hashizume H, Nakagawa S,
Kunitoki K, Sassa Y, Kawashima R (2014) Regional gray matter density is associated with achievement motivation: Evidence
from voxel-based morphometry. Brain Structure and Function 219: 71-83.
Tomoda A, Mizuno K, Murayama N, Joudoi T, Igasaki T, Miyazaki M, Miike T (2007) Event-related potentials in Japanesechildhood chronic fatigue syndrome (CCFS). Journal of Pediatric Neurology 5: 199-208.
Yoshida M, Tanaka M, Mizuno K, Ishii A. Nozaki K, Urakawa A, Cho Y, Kataoka Y, Watanabe Y (2008) Factors influencing theacademic motivation of individual college students. International Journal of Neuroscience 118: 1400-1411.

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Poster Presentations

Reconstruction of the brain from skull fossil using
computational anatomy
Kunihiro Hasegawa1*, Takanori Kochiyama2, Hiroki C. Tanabe1 andNaomichi Ogihara3

1. Graduate School of Environmental Studies, Nagoya University, Japan; 2. Department of Cognitive Neuroscience, Brain Information
Communication Research Laboratory Group, Advanced Telecommunications Research Institute International, Japan; 3. Department of
Mechanical Engineering, Keio University, Japan. *Email: hasegawa.kunihiro@f.mbox.nagoya-u.ac.jp

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176

We investigated the presumed differences in learning abilities between Neanderthals and Modern Humans by
combining evidence from the morphological analysis of fossil brains and the functional mapping of Modern
Humans’ brain functions. Our study faced a fundamental challenge: although we could easily access the Modern Human brain data, the only available data for the Neanderthal’s brain were skull or endocast image reconstructed from fossilized skull produced by Project C01. Hence, to achieve our purpose, we had to establish
a method for extrapolating to human brain functions from skull or endocast anatomy, which is the only data
available for both Modern Humans and Neanderthals. In our previous study, we developed a skull or endocast-based image registration method using the computational anatomy, which is a standard method such asfunctional MRI (fMRI) and voxel based morphometry (VBM) used in neuroimaging research (Kochiyama
et al. 2014; Tanabe et al. 2014). This technique allows us to not only extrapolate the Neanderthal’s brain that
does not exist, but also to investigate the Neanderthal’s brain morphology on the standardized stereotaxic
space that is commonly used in the brain mapping community. Thus, the present study aimed at revealing amorphological difference between Neanderthals and Modern Humans. We quantitatively compare differencesin patterns of regional anatomy using the multivariate analysis of deformation based morphometry (DBM)
and automated anatomical labelling (AAL) parcellation-based morphometry.
We used three fossil skull CT images of Amud 1, Gibralar and La Chapelle-aux-Saints and 512 MR imagesof Japanese Modern Humans. All MRI images were acquired using a 3-Tesla MR scanner (Siemens, Verio)at the National Institute for Physiological Sciences (NIPS). A T1-weighted fine structural whole-brain image were acquired using a magnetization prepared rapid acquisition gradient echo (MP-RAGE). All of thesubjects gave written informed consent for participation in the study. The protocol was approved by the local
medical ethics committee at NIPS. Image processing was performed using the Statistical Parametric Mapping package (SPM8; Wellcome Department of Cognitive Neurology, London, UK), Surfstat (www.math.
mcgill.ca/keith/surfstat/) and in-house software implemented in MATLAB (Mathworks Inc., Sherborn,
MA, USA). First, we create endocast image as follows: For Neanderthal’s data, the reconstruction of fossil
endocast has already been explained in detail (Amano et al. 2014; Ogihara et al. 2014), so we will only briefly
describe them. The fossil skull CT image was first isolated cranial fragments and then they were reassembled
into the correct position. After approximated the cranial surface, the endocast was extracted. For Modern
Human’s data, we segmented individual MR images into the tissue probability maps of GM (gray matter),
WM (white matter) and CSF (cerebrospinal fluid) by segmentation-normalization algorithm in SPM. Then,the sum of these segments image was threshold at the probability of 90 % to create the endocast image. The
residual artifact cleaning with the removal of small isolated segments and filling holes was also applied tocorrect topological errors. Second, these endocasts were enter into Diffeomorphic Anatomical Registration
Through Exponentiated Lie algebra or DARTEL registration algorithm to estimate a globally one-to-one
smooth and continuous mapping between individuals well as between an individual image and a data specifictemplate image. Resulting spatial transformation parameters describe the deformation field that encodeswhole information relating to the position, size, and shape of the brain. Based on DARTEL, we conducted
the quantitative comparison of brain morphology between Neanderthals and Modern Humans using two dif-

Poster Presentations

ferent methodologies. (1) DBM is one of the multivariate analysis techniques, which is useful to detect structural differences over the entire brain because it analyzes positional differences between every voxel and areference endocast. The Hotelling’s T2 test can evaluate differences in the displacement vectors (i.e. the shapeof endocast) between populations. (2) AAL (automated anatomical labeling) parcellation based morphometry(tentatively-named as PBM) was newly developed technique, which can evaluate the difference in regional
brain volume over 116 brain regions defined by AAL. AAL scheme is an anatomical parcellation of the spatially normalized single-subject high-resolution T1 volume (Collin 27 brain) provided by the Montreal Neurological Institute (MNI). To calculate the brain volume in each of 116 regions for each individual, AAL map
was inverse-transformed to each individual participant’s native space by DARTEL transformation. The two
sample T test was performed for each AAL region to evaluate the regional anatomical difference between
populations.
Fig. 1 shows the result of multivariate test (left) for differences in the displacement vectors (right). The comparison of endocast between 512 Modern Humans and 3 Neanderthals showed a significant cluster spanningalmost the entire posterior brain regions, including superior, inferior and posterior parietal cortices, occipitalcortex and cerebellum. Small cluster was found in frontal and limbic regions. The small arrows in Fig. 1 (b)indicate 3D displacements needed to locally deform the Neanderthal’s endocast to match it to the Modern
Human’s endocast. The arrows in cerebellum are long and inward mean large shrinkage in the Neanderthal’sendocast relative to the Modern Humans. The shape change also occurred at the visual and frontal expansion
and the parietal and temporal shrinkage.
Fig. 2 shows the result of the PBM. The left and right row indicates the comparison of AAL-defined volumesof interest between Modern Humans vs. Neanderthals (left), and Neanderthals vs. Modern Humans (right),
respectively. Compared to Modern Humans, Neanderthals showed volume reductions of endocast in widespread areas of both parietal and cerebellum. The area of volume reduction was found in the lateral cerebellum, superior and inferior parietal lobule including angular and supramarginal gyrus. Neanderthals showedvolume expansions compared with Modern Humans, mainly in the visual cortex. Volume enlargements werealso found in the orbitofrontal cortices and limbic structures such as hippocampus. Visual inspection of Figs.
2 and 3 shows the similar local volume reduction/expansion patterns using a different methodology.
In this study, we found that the regional morphological difference between Neanderthals and Modern
Humans in both the cerebral cortex and cerebellum using the computational anatomy approach. In future research, we will explore the structural-functional relations using a combination of the current approach alongwith the meta-analysis about social learning skills and creativity. Through ongoing analysis we hope to uncover the neuro-cognitive functions responsible for Neanderthal-Homo sapiens replacement drama.
References
Amano H, Morita Y, Nagano H, Kondo O, Suzuki H, Nakatsukasa M, Ogihara N (2014) Statistical interpolation of missing
parts in human crania using regularized multivariate linear regression analysis. In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 2: Cognitive and physical perspectives. Springer,
New York, pp. 161-169.
Kochiyama T, Tanabe HC, Ogihara N (2014) Reconstruction of the brain from skull fossils using computational anatomy. In:
Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 2:
Cognitive and physical perspectives. Springer, New York, pp. 191-220.
Ogihara N, Morita Y, Amano H, Kondo O, Suzuki H, Nakatsukasa M (2014) Application of sliding landmark method for morphological analysis of modern Japanese neurocranial shape. In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics of learning in Neanderthals and Modern Humans, vol. 2: Cognitive and physical perspectives. Springer, New York, pp. 145152.
Tanabe HC, Kochiyama T, Ogihara N, Sadato N (2014) Integrated analytical scheme for comparing Neanderthal’s brain toModern Human brain using neuroimaging techniques. In: Akazawa T, Ogihara N, Tanabe HC, Terashima H (eds) Dynamics oflearning in Neanderthals and Modern Humans, vol. 2: Cognitive and physical perspectives. Springer, New York, pp. 203-207.

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Fig. 1 Result of DBM analysis.

Fig. 2 Result of DBM analysis.

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Author Index
Abe-Ouchi, Ayako
Akazawa, Takeru
Akiyama, Hiroyuki
Amano, Hideki
Aoki, Kenichi
Araki, Tsuyoshi
Arai, Saiji
Aripdjanov, Otabek
Barkai, Ran
Bar-Yosef, Ofer
Barzilai, Omry
Belfer-Cohen, Anna
Berna, Francesco
Biagi, Paolo
Biglari, Fereidoun
Bruner, Emiliano
Centi, Laura
Chan, Wing-Le
Chazan, Michael
Condemi, Silvana
Coolidge, Frederick L.
Cortés Sanchez, MiguelDouka, Katerina
Enquist, Magnus
Feldman, Marcus
Finlayson, Clive
Fogarty, Laurel
Furuichi, Takeshi
García-Alix, Antonio
Gasparyan, Boris
Ghirlanda, Stefano
Goring-Morris, Nigel
Hagino, Izumi
Hanawa, SugikoHasegawa, Kunihiro
Hayakawa, Toshiyuki
Hershkovitz, Israel
Hewlett, Barry S.
Hewlett, Bonnie
Higham, Tom
Higurashi, Yasuo
Horiuchi, Shiro

60, 156, 157
18, 131
142
116, 11996, 99109
73
73, 12829125
44
43, 81
26
71
48
112
39
60, 157
26
19
21
159
64
95
96
159
96
93
159
45
95
43
143
109
168, 176
148
38, 4489
88
64, 65135
97

Horwitz, Liora Kolska
26
Hoshino, Yukinobu
171
Hovers, Erella
81
Imamura, Kaoru
142
Ishida, Hajime
116, 119, 148
Ismailova, Jannat
73
Jimenez-Espejo, Francisco J. 159
Jinam, Timothy A.
77
Jöris, Olaf
80
Kadowaki, Seiji
68, 154
Kamei, Nobutaka
90
Kandel, Andrew W.
40, 45
Kanjou, Yousef
131
Kanzawa-Kiriyama, Hideaki 77
Kato, Hirofumi
75
Kawaguchi, Akira
148
Kawahata, Hodaka
59, 156
Kawashima, Ryuta
109
Khenzkhenova, Fedora
75
Kimura, Ryosuke
145, 148
Kobayashi, Yasushi
113, 161
Kobayashi, Yutaka
154
Kochiyama, Takanori
121, 176
Koops, Kathelijne
93
Kondo, Osamu
116, 119, 161, 164, 166
Kondo, Yasuhisa
56, 61
Kosuda, Shigeru
113
Kotozaki, Yuka
109
Koyama, Tadashi
139
Kubo, Daisuke
116, 119, 164
Kunikida, Dai
75
Le Brun-Ricalens, Foni
84
Lind, Johan
95
Lipnina, Ekaterina A.
75
Marder, Ofer
44
Matsui, Toshiyasu
113
Medvedev, German I.
75
Michikawa, Takashi
119, 161, 166
Miki, Takehiro
73
Mitani, Keita
171
Miura, Naoki
168, 171
Mizuno, Kei
174

Monge Gómez, Guadalupe
Morgan, David
Moriguchi, Masaki
Morita, Yusuke
Murayama, Sadayuki
Nagai, Kenji
Naganuma, Masaki
Nakahashi, Wataru
Nakata, Hiroto
Nakatsukasa, Masato
Nishiaki, Yoshihiro
Nouchi, Rui
Nozawa, Takayuki
Obrochta, Stephen P.
Ogihara, Naomichi
Oguchi, Takashi
O’ishi Ryouta
Omori, Takayuki
Omura, Keiichi
Otte, Marcel
Ponce de León, Marcia S.
Prevost, Marion
Rodriguez-Vidal, Joaquin
Ronen, Avraham
Ryu, Heungjin
Sadato, Norihiro
Saitou, Naruya
Sakamaki, Tetsuya
Sano, KatsuhiroSanz, Crickette
Sasaki, Yukako
Sato, Takao
Sato, Takehiro
Sharon, Gonen
Shemer, Maayan
Shidrang, Sonia
Shimelmitz, Ron
Song, WonsuhStarnini, Elisabetta
Sugiura, Motoaki
Suleymanov, Rustam
Suzuki, Kenji

159
93119, 161, 166
119
116
171
154
145, 151
73, 133
119
73, 124, 131
109
109
59
116, 119, 121, 161, 164, 166, 176
56, 156
60, 157
66, 67, 73
108
23
115
39
159
31
93
121
77
93
66, 67
93
168
75
148
34
39
51
40
56
71
109
73
75

Suzuki, Hiromasa
Takakura, Jun
Takahashi, Kunio
Taller, Andreas
Tamura, Kohei
Tanabe, Hiroki C.
Terashima, Hideaki
Tokuyama, Nahoko
Tostevin, Gilbert B.
Tsatskin, Alexander
Varoner, Oz
Wakano, Joe Yuichiro
Walker, Steven
Watanabe, Chiaki
Watts, Ian
Weinstein-Evron, Mina
Wilkins, Jayne
Wynn, Thomas
Yamagami, Eiko
Yamaguchi, Kyoko
Yamamoto, Ken
Yamauchi, Taro
Yeshurun, Reuven
Yogi, Akira
Yokoyama, Ryoichi
Yokoyama, Yusuke
Yoneda, Minoru
Yoshida, Kunio
Zaidner, Yossi
Zollikofer, Christoph P. E.

119, 161, 164, 166
85
157
45
145
116,121,168,171,176
106
93
82
38
39
96, 102
26
148
26
38
26
21
137
148
148
36, 143
38
116
109
59, 156
156
75
38, 39115

RNMH2014
The Second International Conference November 30 – December 6, 2014, Date City
Edited by Takeru Akazawa & Yoshihiro Nishiaki

RNMH PROJECT SERIES No. 003

RNMH2014
The Second International Conference

Replacement of Neanderthals by Modern Humans:
Testing Evolutionary Models of Learning

November 30 December 6, 2014, Date City

Program and Abstracts
Edited by

Takeru Akazawa &Yoshihiro Nishiaki