RNMH2012
The First International Conference November 18-24, 2012, Tokyo
Edited by Takeru Akazawa & Yoshihiro Nishiaki

RNMH PROJECT SERIES No. 002

RNMH2012
The First International Conference

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

November 18-24, 2012, Tokyo
Edited by

Takeru Akazawa &Yoshihiro Nishiaki

RNMH2012
The First International Conference

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

November 18-24, 2012, Tokyo
Edited by

Takeru Akazawa &Yoshihiro Nishiaki

Editor-in-Chief

Takeru Akazawa

Research Institute, Kochi University of Technology

Editorial Board

Shunichi Amari

Brain Science InstituteKenichi 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
Nagoya University

Hideaki Terashima
Kobe Gakuin University

Minoru Yoneda

The University of Tokyo

RNMH Project Series

Takeru Akazawa and Yoshihiro Nishiaki (eds.) 2012
RNMH2012 ―The First International Conference on the
Replacement of Neanderthals by Modern Humans: Testing
Evolutionary Models of Learning

Published by

RNMH Project Group

Research Institute, Kochi University of Technology
CIC Tokyo, 3-3-6 Shibaura, Minato-ku, Tokyo 108-0023, Japan
Te l: +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 Research on Innovative Areas(Grant No. 22101001) from Japanese Ministry of Education,
Science, Culture, and Technology.
ISBN: 978-4-9906371-0-1© 2012 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, orotherwise, without the prior permission of the RNMH projectgroup.

Produced by Akita Kappan, Inc., Akita
Designed by Tsukasa FujitaPreface

006

Conference Program			

009

			

Oral Presentations

The Opening Keynote Lectures

024

The Process of RNMH: Global and Comprehensive
Explanatory Models

034

Learning Behaviors in Prehistoric and Modern
Humans045

Human-Specific Learning Strategies and Cultural
Evolution

056

Towards Digital Reconstruction of Fossil Crania andtheir Brain Morphology 		

066

Neuroimaging Approach to the Learning		Hypothesis		

076

Poster Presentations

Group A01: Palaeolithic Archaeology			 086Group A02: Hunter-Gatherer Studies

		 097

Group B01: Evolutionary Models			 121Group B02: Climate and Geoscience			 137Group C01: Skull and Brain				 151Group C02: Neuroscience			

161

Author Index

167

C o n t e n t sModern humans, whose origins lie in the African

continent of around 200,000 years ago, began tomigrate to the Middle East and Europe by 50,000
years ago, replacing the Neanderthals who had

originally inhabited those regions. What actually
happened when modern humans encountered
Neanderthals? There may have been someinterbreeding, but ultimately modern humans wereto replace Neanderthals. The background to theextinction of the Neanderthals, anatomically distinct
from modern humans while still being extremely

close to them, and the contrasting success of modern
humans is a research topic of unmatched interest in

the debate surrounding the emergence and spread ofmodern humans.

There are already a number of theories for the

P r e f a c e Replacement of Neanderthals by Modern Humans(RNMH), including the neural theory and the
population theory. What many of the existing

theories have in common is the suggestion thatthe cultural and technological adaptive capacities
of Neanderthals and modern humans to theirenvironments were different. The adaptability ofhumans is heavily dependent upon culture, andculture is fostered through learning. From this, it iseasy to imagine how the nature of learning could
have been a major factor for the divergence in theadaptive capacities of the two populations. Was thedifference between the relative learning capacities of
Neanderthals and modern humans inherent? And

was it a decisive factor in causing the replacement of
Neanderthals by modern humans? This supposition
has been deemed the “learning hypothesis,” and006the RNMH Project was launched in 2010 in order

be given by researchers working in fields that haveapproaches.

prehistoric archaeology, theoretical biology, cognitive

to examine this working hypothesis from multiple
The Project brings together around 80 researchers

from multiple fields, engaged in collaborative

research. Prehistoric archaeology examines what

physical evidence there is to support the differencesin the learning behaviors of the two populations,while cultural anthropology looks at the natureof learning in modern hunter-gatherer culturesand creates interpretive models for archeologicalrecords. Theoretical biology provides theoreticalevidence supporting the learning hypothesis,
while paleoclimatology and environmental science

made significant contributions to this field so far—archaeology, fossil engineering—followed by fivesessions on core themes: (1) The process of RNMH:

global and comprehensive explanatory models,(2) Learning behaviors in prehistoric and modern

humans, (3) Human-specific learning strategies andcultural evolution, (4) Towards digital reconstructionof fossil crania and their brain morphology, and (5)Neuroimaging approach to the learning hypothesis.
The Conference will also feature a large number ofrelated poster presentations.

Over six days, the Conference will be a platform forprovide supporting evidence for the differences inlively debate that will hopefully lead to progress in ouradapt to environments. Fossil engineering is used toand that of us, modern humans. Of course, thethe respective capabilities of both populations tounderstand the morphology of the brains of fossil
hominids, while neuroscience seeks to map the partsof the brain related to learning.

The Project is initially being conducted on a five-

year basis. This Conference has been planned tomark the mid-way point of the Project, and to set
out the direction and goals of the Project going

forward. This Conference has two specific objectives:
First, the presentation, by a number of Japanese
researchers and joint researchers from overseas, ofinterim reports detailing progress in the RNMH

understanding of the true nature of the Neanderthals
verification of the learning hypothesis is our goal, but

a greater achievement would be the simultaneous
generation of a new problem-solving perspective. The
Conference has been generously supported by the
Ministry of Education, Culture, Sports, Science andTechnology of Japan. Thanks are also extended to themembers of the Steering Committee for Scientific

Research on Innovative Areas, as well as staff at the
National Institute of Informatics, who organized thevenue.

Project. Second, the measurement of progress in

Takeru Akazawa

by having integrated the successes achieved thus

Yoshihiro Nishiaki

verifying and developing the learning hypothesis
far by other researchers working on similar research

Kochi University of Technology
The University of Tokyo

projects in related fields. Keynote presentations will

007

RNMH2012

The First International Conference

Kenichi Aoki

Department of Biological Sciences
Graduate School of Sciences,The University of Tokyo
7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
Tel: +81-(0)3-5841-4485 Fax: +81-(0)3-3818-7547
kenaoki@biol.s.u-tokyo.ac.jp

Minoru Yoneda

ORGANIZING COMMITTEE
Takeru Akazawa

Research Institute, Kochi University of Technology
3-3-6 Shibaura, Minato-ku, Tokyo 108-0023, Japan
Tel: +81-(0)3-5440-9039 Fax: +81-(0)3-5440-9119
akazawa.takeru@kochi-tech.ac.jp

Yoshihiro Nishiaki

The University Museum, The University of Tokyo
7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
Tel: +81-(0)3-5841-2491 Fax: +81-(0)3-5841-8451
nishiaki@um.u-tokyo.ac.jp

Hideaki Terashima

Faculty of Humanities and Sciences
Kobe Gakuin University
518 Arise, Ikawadani-cho, Nishi-ku, Kobe 651-2180, Japan
Tel: +81-(0)78-974-1551 Fax: +81-(0)78-976-7296
terasima@human.kobegakuin.ac.jp

Head Office RNMH

008

Research Institute, Kochi University of Technology
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

The University Museum, The University of Tokyo
7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
Tel: +81-(0)3-5841-2483 Fax: +81-(0)3-5841-8451
myoneda@um.u-tokyo.ac.jp

Naomichi Ogihara

Department of Mechanical Engineering, Keio University
3-14-1 Hiyoshi, Kohoku-ku, Yokohama 223-8522, Japan
Tel: +81-(0)45-566-1423 Fax: +81-(0)45-566-1495
ogihara@mech.keio.ac.jp

Hiroki C. Tanabe

Graduate School of Environmental Studies
Nagoya University
Furo-cho, Chikusa-ku, Nagaoya, Aichi 464-8601, Japan
Tel: +81-(0)52-789-2256 Fax: +81-(0)52-789-2221
htanabe@nips.ac.jp

Shunichi Amari
Shiro Ishii
Tasuku Kimura
Ofer Bar-Yosef
Ralph L. Holloway

Conference Program
Oral Presentations

Each presentation consists of a 40-minute talk, followed by a 15-minute discussion.

Sunday, November 18, 2012The Opening Keynote Lectures
12:00–13:00

Registration

13:00–13:15

Takeru Akazawa, Kochi University of Technology, Japan				Opening Remarks		

Chair: Yoshihiro Nishiaki
13:15–14:10

Ofer Bar-Yosef, Harvard University, USA						 024Neanderthals and Modern Humans – A Global View: Isolation, Expansion, 		Competition, and Demise

14:10–15:05

Nicole Creanza, Laurel Forgarty, and Marcus W. Feldman*, Stanford University, USA		 027

15:05–15:20

Coffee Break

15:20–16:15

Steven J. Mithen, University of Reading, UK						 028

Exploring Cultural Niche Construction from the Paleolithic to Modern Hunter-Gatherers

Neanderthals and Modern Humans – A Cognitive View: Intelligence, Symbolism
and Language

16:15–17:10

Christoph P. E. Zollikofer* and Marcia Ponce de LeÓn, University of Zurich,
Switzerland

The Reconstitution of the Neanderthal Brain
17:30–

030

Welcome Party009

Oral Presentations
Monday, November 19, 2012

The Process of RNMH: Global and Comprehensive Explanatory
Models
9:00–10:00

Registration

Chair: Seiji Kadowaki
10:00–10:55

João Zilhão, University of Barcelona, Spain034

10:55–11:50

Ran Barkai* and Avi Gopher, Tel Aviv University, Israel

035

The Replacement of Neanderthals by Modern Humans in Europe

Cultural and Biological Transformations in the Middle Pleistocene Levant: A View
from Qesem Cave, Israel

11:50–13:00

Lunch Break

13:00–13:40

Poster Session: Group A01/A02

Chair: Akira Ono
13:45–14:40

Seiji Kadowaki* and Katsuhiro Sano, Nagoya University, Japan

038

From the Beginning: Tracing the Patterns of Cultural Changes of Early Homo sapiens in
Africa, West Asia, and Europe

14:40–15:35

Hirofumi Kato* and Masaki Naganuma, Hokkaido University, Japan

15:35–16:00

Coffee BreakThinking the Beginning of Early Upper Palaeolithic in Siberia and Central Asia

040

Chair: Naoko Matsumoto
16:00–16:55

Ayako Abe-Ouchi*, Yusuke Yokoyama, Hodaka Kawahata, Takashi Oguchi,

and Minoru Yoneda, The University of Tokyo, Japan					 042Investigation of Past Climate and Vegetation for Understanding the Replacement of
Neanderthals by Modern Humans

16:55–17:50

Steven L. Kuhn, University of Arizona, USA 						Cultural Transmission, Institutional Continuity and the Persistence of the Mousterian

Closing Remarks

010

044

Oral Presentations
Tuesday, November 20, 2012

Learning Behaviors in Prehistoric and Modern Humans
9:00–10:00

Registration

Chair: Hideaki Terashima

10:00–10:55

Hideaki Terashima, Kobe Gakuin University, Japan					 045

10:55–11:50

Jun Takakura, Hokkaido University, Japan 						 048

Evolutionary Development of Learning and Teaching Strategies in Human Societies

New Insights into the Skill Learning Processes in the Lithic Production: An Analysis of the
Refitted Material from the Kyushirataki 15 Site in Hokkaido, Northern Japan

11:50–13:00

Lunch Break

13:00–13:40

Poster Session: Group A01/A02

Chair: Steven L. Kuhn
13:45–14:40

Yoshihiro Nishiaki, The University of Tokyo, Japan 					 050
“Gift” as a Means of Cultural Transmission: An Archaeological Implication of Bow andArrow Technology in Papua New Guinea

14:40–15:35

Shiro Horiuchi, and Sachiko Kubota

15:35–16:00

Coffee Break

16:00–16:55

Barry S. Hewlett, Washington State University, USA 			The Effects of Cross-Boundary Rituals on Cultural Innovation

Teaching, Trust, and Early Rapid Social Learning in Hunter-Gatherers

053

		 055

Closing Remarks

Wednesday, November 21, 2012

Holiday/Excursion

011

Oral Presentations
Thursday, November 22, 2012

Human-Specific Learning Strategies and Cultural Evolution
9:00–10:00

Registration

Chair: Kenichi Aoki
10:00–10:55

Kenichi Aoki, The University of Tokyo, Japan056

10:55–11:50

Joe Yuichiro Wakano, Meiji University, Japan

058

Determinants of Cultural Evolutionary Rates

Scheduling of Individual and Social Learning as Optimal Life History Strategy

11:50–13:00

Lunch Break

13:00–13:40

Poster Session: Group B01/B02/C01/C02

Chair: Laurent Lehmann
13:45–14:40

Laurent Lehmann, University of Lausanne, Switzerland

The Handaxe and the Microscope: Experience-Based and Fictitious Learning in a060

Geometric Model of Adaptation
14:40–15:35

Nobuyuki Takahashi*, Ayaka Hatano, Misato Inaba, Ryoichi Onoda, and Dora Simunovic,
Hokkaido University, Japan

		Social Learning, Trial-and-Error, and Creativity
15:35–16:00

Coffee Break

16:00–16:55

Alex Mesoudi, Durham University, UK

Experimental Studies of Modern Human Social and Individual Learning in an
Archaeological Context: People Behave Adaptively, but within Limits

Closing Remarks

012

061

063

Oral Presentations
Friday, November 23, 2012

Towards Digital Reconstruction of Fossil Crania and their Brain
Morphology
9:00–10:00

Registration

Chair: Naomichi Ogihara

10:00–10:55

Naomichi Ogihara*, Takeo Kikuchi, Yusuke Morita, Hideki Amano, Osamu Kondo,
Masato Nakatsukasa, Hiromasa Suzuki, Takashi Michikawa, Hajime Ishida,
and Takeru Akazawa, Keio University, Japan

066

Emiliano Bruner, Centro Nacional de Investigación sobre la Evolución Humana, Spain

068

Digital Reconstruction of the Neanderthal Amud 1 Cranium
10:55–11:50

Functional Craniology, Human Evolution, and Anatomical Constraints in the Neanderthal
Braincase

11:50–13:00

Lunch Break

13:00–13:40

Poster Session: Group B01/B02/C01/C02

Chair: Marcia Ponce de Leon
13:45–14:40

Osamu Kondo*, Daisuke Kubo, and Naomichi Ogihara, The University of Tokyo, Japan

14:40–15:35

Yasushi Kobayashi*, Toshiyasu Matsui, Yoshinori Haizuka, Naomichi Ogihara,

Virtual Endocast of Qafzeh 9, A Representative of Early Modern Humans

Naoki Hirai, and George Matsumura, National Defense Medical College, Japan
Cerebral Sulci and Gyri Observed on Macaque Endocasts

15:35–16:00

Coffee Break

16:00–16:55

Takanori Kochiyama*, Hiroki C. Tanabe, and Naomichi Ogihara, Kyoto University,
Japan

Reconstruction of the Brain from Skull Fossil Using Computational Anatomy

070

072

074

Closing Remarks013

Oral Presentations
Saturday, November 24, 2012

Neuroimaging Approach to the Learning Hypothesis: Learning,
Creativity, and Social Cognition
Chair: Hiroki C. Tanabe
9:00–9:55

Hiroki C. Tanabe, Nagoya University, Japan076

9:55–10:50

Dietrich Stout, Emory University, USA

078

10:50–11:10

Coffee BreakInnovative Society and Social Brains

Social Neuroscience of Stone Tool-Making Skill Acquisition

Chair: Dietrich Stout
11:10–12:00

Hiroaki Kawamichi*, Kazufumi Yoshihara, Ryo Kitada, Masahiro Matsunaga,

Akihiro Sasaki, Yumiko Yoshida, Haruka Takahashi, and Norihiro Sadato, National Institutefor Physiological Sciences, Japan079

Kei Mizuno, RIKEN Center for Molecular Imaging Science, Japan

082

Study Implication Related to Sense of Acceptance: Key Factor of Social Learning
12:00–12:55

Neural Substrates of Motivation to Learn and Fatigue

Closing Remarks
13:15–

014

Sayonara Party

Conference Program
Poster Presentations

Note for the authors: The board available for each poster presentation is 90 cm wide and 210 cm high. Pleasesubmit your poster at the registration desk, and present it at the assigned core time. We request the authors toremove your poster before 17:00 on November 23.

Group A01: Palaeolithic Archaeology core time on November 19 & 20
(1) Yasuo Higurashi, Osaka University, Japan

Aimed Spear-Throwing in Modern Humans: A Preliminary Study of Upper Limb Kinematics

		(2) Yasuhisa Kondo, Tokyo Institute of Technology, Japan

086

088

Neander DB 2.0: Improvement of the “Replacement of Neanderthals by Modern Humans” Lithic
Industry Database(3) Hiroto Nakata, Aoyama Gakuin University, Japan090(4) Yoshihiro Nishiaki, The University of Tokyo, Japan

092(5) Jun Takakura, Hokkaido University, Japan

094

The Initial and Early Upper Paleolithic of the Japanese Archipelago

The Archaeological Database for the RNMH Project

Lithic Refitted Materials and Identification of Technical Skill Level: Lessons from the Upper
Paleolithic Assemblages of the Shirataki Sites in Hokkaido, Northern Japan

015

Poster Presentations
Group A02: Hunter-Gatherer Studies core time on November 19 & 20
(6) Juko Ando, Keio University, Japan								 097Comparison between Individual, Imitative and Instructive Learning(7) Adam Howell Boyette, Washington State University, USA099

Elements of Social Learning among Hunter-Gatherer and Farmer Children in the Central African
Republic(8) Izumi Hagino* and Taro Yamauchi, Hokkaido University, Japan

Daily Physical Activity and Time-Space Allocation of Pygmy Hunter-Gatherers’ Children in

101

Southeast Cameroon(9) Koji Hayashi*, Ryota Ishii, Yuuki Nakamura, Hideaki Terashima, and Yoshihiro Nishiaki,
Kobe-Gakuin University, Japan

103

Technical Transmission of Hunting Tool Manufacture: A Case of Spear Hunting Among Modern
Hunter-Gatherers in Southeast Cameroon(10) Bonnie Hewlett, Washington State University, USA“Finding the Spirit to Create” Innovation and Social Learning Among Aka Adolescents of the105

Central African Rainforest(11) Kaoru Imamura, Nagoya Gakuin University, Japan
Hunting Play among San Children

016

108

(12) Nobutaka Kamei, Aichi Prefectural University, Japan

110

Play Activities and Cultural Transmission among Hunter-Gatherer Children: A Case of the Baka in
Cameroon(13) Tadashi Koyama, Kobe Gakuin University, Japan

Cognitive Flexibility and Making Objects in Baka Children

112(14) Keiichi Omura, Osaka University, Japan							 114The Art of Patience as the Emotional Basis for Creativity and Cultural Learning: Learning andEcological Adaptation among the Canadian Inuit(15) Akira Takada, Kyoto University, Japan								 116Communicative Musicality and Learning in Caregiver-Child Interactions among the San of NorthCentral Namibia(16) Eiko Yamagami, Kobe Gakuin University, JapanThe Resilience of Pygmy Children– On Their Drawings about Favorite Things

117(17) Taro Yamauchi* and Izumi Hagino, Hokkaido University, Japan 					 119Estimation of the Period of Childhood and Child Growth Characteristics of Pygmy Hunter-		Gatherers in Southeast Cameroon

017

Poster Presentations
Group B01: Evolutionary Models core time on November 22 & 23
(18) Shiro Horiuchi, Shibaura Institute of Technology, Japan

121(19) Kohkichi Kawasaki, Doshisha University, Japan

123(20) Ryosuke Kimura, University of the Ryukyus, Japan125(21) Yutaka Kobayashi, The University of Tokyo, Japan127

The Effects of Cross-Boundary Rituals on Cultural Diversity

Effect of a Heterogeneous Environment on Range Expansion of Individual and Social Learners

Dispersals of Modern Humans: Inferences from Genome Diversity Data

A Simulation Study on the Replacement of Neanderthals by Modern Humans in Europe: 		Implications of Climate Change, Cultural Diversification, and the Shape of the Continent(22) Wataru Nakahashi, Meiji University, Japan							 130A Mathematical Model of Cultural Interactions between Modern and Archaic Humans(23) Makoto Shimada, Fujita Health University, JapanAn Attempt to Estimate the Functional Difference in Haplotypes between Archaic and Modern

133

Humans(24) Lucas Henriques Viscardi*, Vanessa R. Paixão- CÔrtes, Francisco Mauro Salzano, Tábita Hünemeier,and Maria Cátira Bortolini, Universidade Fedoral do Rio Grande - Institute de Ciencias Humanas eda Informação, Brazil

The Down of Hierarchy: The Cognitive Ability of Archaic Hominins

018

135

Poster Presentations
Group B02: Climate and Geoscience core time on November 22 & 23
(25) Wing-Le Chan*, Ayako Abe-Ouchi, Ryota O'ishi, and Kunio Takahashi,The University of Tokyo,
Japan

137

Using Numerical Models to Simulate Climate and Vegetation Changes during the Last Glacial Period(26) Mark C Diab* and Minoru Yoneda, The University of Tokyo, Japan

Late Middle Palaeolithic (OIS 4/3) Neanderthal Prey Exploitation and Land Use at Dederiyeh

139

Cave, Syria Based on Multiple Isotopic Analyses of Intra-Tooth Enamel and Mortality Profiles ofWild Goat, Red Deer, and Gazelle(27) Yasuhisa Kondo* and Takashi Oguchi, Tokyo Institute of Technology, Japan			 141How can We Apply Ecological Niche Models to Palaeoanthropological Research?

(28) Yuzo Marukawa*, Hirohisa Mori, Yoshifumi Nakamura, and Takeru Akazawa, International ResearchCenter for Japanese Studies, Japan

Bibliographic Database for “Replacement of Neanderthals by Modern Humans”

143(29) Stephen Obrochta*, Yusuke Yokoyama, and Hodaka Kawahata, The University of Tokyo, Japan145(30) Takayuki Omori* and Minoru Yoneda, The University of Tokyo, Japan147(31) Song Wonsuh*, Yasuhisa Kondo, and Takashi Oguchi, The University of Tokyo, Japan

149

Plant Creating a Common Chronology for Levant Pollen Records Based on Ice Core Synchronization

Population Dynamics and Spatial Distribution Inferred from Radiocarbon Meta Analysis

PaleoGeo: A Web GIS for Distributing Information from Paleoenvironmental Literature

019

Poster Presentations
Group C01: Skull and Brain core time on November 22 & 23(32) Hideki Amano*, Yusuke Morita, Hiroyasu Nagano, Osamu Kondo, Hiromasa Suzuki,
Masato Nakatsukasa, and Naomichi Ogihara, Keio University, Japan
Statistical Interpolation of Missing Parts in Fossil Crania

151(33) Hiroyuki Hishida*, Hiromasa Suzuki, Takashi Michikawa, Yutaka Ohtake, Satoshi Oota, Naomichi
Ogihara, and Osamu Kondo, The University of Tokyo, JapanCT Image Segmentation Using FEM with Optimized Boundary Condition(34) Daisuke Kubo* and Hiroki C. Tanabe, The University of Tokyo, Japan
Cerebellar Size Estimation from Endocranial Measurements

153

155

(35) Masaki Moriguchi*, Hiromasa Suzuki, Takashi Michikawa, Naomichi Ogihara, and Osamu Kondo,
Chuo University, Japan

Transferring Semi-Landmarks on Surfaces(36) Yusuke Morita*, Hideki Amano, Masato Nakatsukasa, Osamu Kondo, and Naomichi Ogihara,
Keio University, Japan

A Geometric Morphometric Study of Neurocranial Shape Variations in the Crania of Modern
Japanese

020

157

159

Poster Presentations
Group C02: Neuroscience core time on November 22 & 23(37) Yukinobu Hoshino*, Naoki Miura, and Kenji Nagai, Kochi University of Technology, Japan
Motion Analysis for Stone-Knapping of the Skilled Levallois Technique

161(38) Naoki Miura*, Kenji Nagai, Mika Yamazaki, Yumiko Yoshida, Hiroki C. Tanabe, Takeru Akazawa,
and Norihiro Sadato, Tohoku Institute of Technology, Japan

Brain Activation Related to the Imitative Learning of Bodily Actions Observed during the 		164

Construction of a Mousterian Stone Tool: A Functional Magnetic Resonance Imaging Study

021

Keynote
Oral Presentations

The Opening Keynote

Lectures

RNMH2012

Neanderthals and Modern Humans – A Global View:
Isolation, Expansion, Competition and DemiseOfer Bar-Yosef

Department of Anthropology, Harvard University, USA

The Opening Keynote Lectures

024

Neanderthals, a European population that emergedsome 400-200,000 years ago (or earlier around600-400,000 years) was undoubtedly successful insurviving through several glacial periods. The remainsof their past life ways include human fossils (buried ornot), food refuse (mostly bones, rare plants), preserved
hearths in particular climatic conditions (such as
the Mediterranean basin), and often plenty of stone
tools. The information, published in more than one
language, was collected through the excavations of
numerous sites since the mid-Nineteen century. The
Neanderthals are generally considered as Europeans
who lived through the period we call “Middle
Paleolithic”, a relative chronological term formed
prior to the time when radiometric techniques fordating were not available.
The Neanderthals possibly lived in small
communities that were spread over large territories
but maintained their relationships in order to keep
their mating systems and secure their biologicalsurvival. When viewed through the few publishedsamples of ancient DNA or their teeth across Eurasia,
it seems that they represent a particular population.
Morphological deviations from the western European
Neanderthals were generally found at the edges of
their spatial distribution.
Neanderthals emerged from what
paleoanthropologists call Homo heidelbergensis andfollowing their expansions into the Mediterraneanbasin as well as further east, reaching the Altai
mountains, they interacted and mixed with otherpeople, perhaps the direct descendants of the Asian
Homo erectus who inhabited these lands in earlier times.

We often identify the Middle Paleolithic stone
tool assemblages based on technological (various
core reduction techniques) and typological aspects(i.e., blanks that were shaped into tools). Amongthe latter archaeologists define side scrapers andpoints, shaped by retouch, some of which wereconstantly resharpened. Special types are large andsmall handaxes (considered as indicating culturalcontinuation from the earlier Acheulian Complex),and foliates. Their assemblages fall mainly under the
categories of Mousterian of Acheulian Tradition
and Eastern Micoquian. Using technological andtypological traits, the Mousterian Complex wasclassified as a large entity by the mid-twentiethcentury, and one can geographically trace the
Neanderthal “cultures” within this complex. Each
recognized prehistoric “culture” is based on theassumption that the tool-kits are the technicalexpressions of people who kept manufacturing theirtraditional artifacts regardless of environmental
fluctuations or even when they moved to a new
area. For example, we can identify the bearers of two
different stone tool making traditions, namely, theEastern Micoquian with the foliates and anotherwith Levallois convergent points, who camped in
the same area of the Altai mountains. The evidence
for their presence there was demonstrated by thediscoveries of lithics, fossils and ancient DNA.
However, the recent surprise brought by this typeof biological analysis in the Altai was the discoveryof an unknown population called the Denisovans.
Thus, if we assume that the correlation between
the fossils and the lithic industries is rather simple,

known from the European plains. Further east
the Neanderthals are found in Uzbekistan, Siberia
and their industries near the Yellow River. It ishypothesized that Neanderthal remains (or perhaps
the Denisovans) should be expected in northern
China. Therefore, in a few cases we already recognize
territorial boundaries of Neanderthal groups(cultures? tribes?) across Eurasia.
The appearance of the modern humans is now
suggested to have occurred around 60-50,000 years
ago and there are no doubts about their African
origins or the few paths they took as they migrated
into Eurasia. We often attribute to modern humans
a series of cultural traits although not all were areal cultural “package” and a few emerged in Africa
during earlier period. These include high degreeof mobility, sings for group identity, and use ofornaments. Thus moderns were capable of symbolic
behavior that is expressed in the Franco-Cantabrianregion by cave art, mobile art objects (also in otherregions), and in a few localities by open-air rock art.
Among their domestic activities scholars often cite
the use of fire and stones for warmth banking, higher
efficiency in hunting, evidence for feasting, and more.
Among the lithics we stress the blade making which
requires much less skill than producing Levallois
products. But we should mention that not all modern
humans made blades as for example the colonizersof Australia some 45,000 years ago who produced
flake tools (while blades arrived there only in the
Holocene). Not all modern humans groups sharedartistic expressions. Similarly shell beads that alreadywere shaped in a few Middle Paleolithic sites, were
not common in all Upper Paleolithic contexts.
In reviewing the interactions between the two
populations we should take into account thecontinental-wide archaeological information
concerning the lithic industries of local Neanderthals.
The best records are available from all over Europe aswell and western Asia.
During the Upper Pleistocene Neanderthals intemperate Europe responded to climatic calamities byshifting and expanding their territories into westernand central Asia. They were undoubtedly successful
survivors as the recently published information

The Opening Keynote Lectures

now, as archaeologists we challenged to uncover theculture of the Denisovans, who biologically emerged,
as the current interpretation suggests, from Homo
heidelbergensis but later than the Neanderthals.
Indeed, we already encountered a similar issue when
the study of Middle Paleolithic fossils in the Levant
took place from the 1930’s onward. The Levantine
Mousterian Complex of this region is currently
divided into three industries, often uncovered in a
stratigraphic order, from about 220/250,000 through50,000 years ago, and are known as “Tabun D-type,C-type and B-type” or as Early, Middle and Late
Levantine Mousterian. Various methods of Levallois
technique characterize the Middle and Late
Mousterian industries. The assemblages of the Late
Levantine Mousterian, rich in Levallois triangularpoints, contained burials and isolated fragments oflocal Neanderthals (e.g., Dederiyeh, Kebara, andAmud caves as well as layer B in Tabun cave). Thesefossils differ slightly in their skull morphology fromthe “classical European Neanderthals”. The main
surprise occurred already in the 1930’s when thefossils uncovered with “Tabun C-type” (Middle
Mousterian) artifacts in well arranged graves in Skhul
and Qafzeh caves, were classified as a type of Archaic
Homo sapiens, once even labeled as “Proto-CroMagnons” and considered until the early 1980’s as
the ancestors of modern humans. Perhaps they were,
but further discussion is beyond the scope of thispresentation.
Hence, the expansion of the Neanderthals into
western Asia is fully supported by the Levantine
fossils (dated to post - 80/70,000 years ago) andinclude those uncovered in Shanidar cave in the
Zagros mountains (northern Iraq) and the skull
fragment found in Sakjia cave, in the southernfoothills of the Caucasus, as well as the human burial
in Mezmaiskaya cave (Russia). The human groups
who occupied the Taurus, Zagros and southern
Caucasus mountain areas made industries rich inretouched pieces (scrapers and points). They differedfrom the sites on the northern slopes of the Caucasus,
represented by the finds from Mezmaiskaya cave,where the tool kits contained the small bifaces or
foliates and were part of the Eastern Micoquian025concerning the Campanian volcanic eruption in
Europe demonstrates. Years ago we noted that during
the cold period of MIS4 (ca. 75-57,000 years ago)
Neanderthals in the north European plain moved
into refugia in southwest and southeast Europe
resulting in the depopulation of a large region in thenorthern latitudes. Hence, contrary to the prevailingview of several western European scholars during thelast decade it was not the worsening conditions of
MIS3 that caused the Neanderthals demise but theactivities of modern humans - the new migrants.
Unfortunately, not all archaeologists andpaleoanthropologists are familiar with the changing
patterns of behavior of hunters-gatherers, or withissues of human reproduction. Foragers have three
options when they encounter newcomers in theirterritory. They can greet them friendly and evenbiologically mix with them, ignore their presence or
try to deter them physically from exploiting theirtraditional resources. Genetic evidence indicates thata certain degree of interbreeding did occur and thus
all recent Eurasian populations bear Neanderthal
genes. None of the African people have them andthus it becomes obvious that the mixing took placein this continent. There is some archaeological

The Opening Keynote Lectures026

evidence to support it in the records of the so-called“transitional industries” (from the Middle to theUpper Paleolithic) although not all are “transitional”
in the sense of cultural continuity. Hence, thelong period of isolation, at least in the European
region of this vast continent, Neanderthals became
gradually extinct. A possible explanation is the effectsof being cut-off the best territories for hunting
and gathering due to their occupation by modern
humans. The taking over certain critical terrains by
the new migrants severed the long distance breeding
networks of the Neanderthals. Reduction of the total
fertility rate, for example, would naturally lead to thedisappearance of “tribal populations”. In addition weshould consider the possibility of physical conflicts.
Thus, Neanderthal groups became extinct some fasterand earlier chronologically than others, while modern
humans thrived.
Reference
Bar-Yosef, O. (2011) A geographic over view of
Neanderthal-modern human encounters. In: Casting the
Net Wide: Papers in Honor of Glynn Isaac and His Approach to
Human Origins Research, edited by J. Sept and D. Pilbeam,
pp. 193-212. American School of Prehistoric Research
Monograph. Oxbow Books, Oxford.

Figure 1. The expansion of the Neanderthals.

Figure 2. The Neanderthal retreat across Europe.

Exploring Cultural Niche Construction from the
Paleolithic to Modern Hunter-Gatherers

Nicole Creanza, Laurel Fogarty, and Marcus W. Feldman
Department of Biological Sciences, Stanford University, USA

Ni c h e c on s t r u c t i on i s a n e vo l u t i on a r y
process through which organisms modify theirenvironments, thereby altering the selection
pressures on future generations. Cultural niche
construction theory concentrates on those changes
made by the spread of socially learned traits orto the evolutionary niche in which these sociallylearned traits evolve. Here we aim to use a modelof cultural niche construction to elucidate thepossible role of culture in the replacement of
Neanderthals by modern humans.
We focus particularly on two important featuresof human populations: cooperation and teaching.
We find that in a cultural niche construction
framework, the presence of teaching enables

beneficial innovations to spread faster and moreeasily, making them more likely to fix or persist
in a population. This may provide a selective
advantage over species unable to teach or whotransmit information with less fidelity thanmodern humans.
Finally, we show that strong, high fidelity
information transmission in early modern humans
could have enabled cooperative hunting and meatsharing practices to persist in a population whereotherwise they would not. We hypothesize thatthe group-level benefits of cooperation and meat
sharing could have given modern humans anadvantage over similar species incapable of thesepractices.

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027

Neanderthals and Modern Humans – A Cognitive
View: Intelligence, Symbolism and Language

Steven J. Mithen
University of Reading, UK

The Opening Keynote Lectures

028

Ever since the skeletal remains of Neanderthals
were first discovered and debate began about theirevolutionary relationship to Homo sapiens, speculation
has occurred about their cognitive and linguistic
abilities. Dramatically contrasting interpretations of
the same archaeological evidence have been forwarded,
positing Neanderthals as either having a radicallydifferent, and for some inferior, form of mentality thanthat of H. sapiens or having precisely the same cognitive
capacity with regard to both intelligence and language.
Those archaeologists who refute any differences incognition invoke contrasts in environment, demography
and/or society to explain any dissimilarity in thearchaeological records of H. neanderthalensis and H.
sapiens and, ultimately, why the latter remains extant
today and the former is extinct.
This debate has recently been enlivened by claims for
new evidence for Neanderthal symbolism, adding tothose which have been long proposed but contested bythose archaeologists who dispute Neanderthals wereable to think and behave in a symbolic manner. In thispaper I provide a critical summary of the debates andchallenge the view that there is indeed a meaningful
opposition between cognitive and environment/
demographic explanations for differences between H.
neanderthalensis and H. sapiens. This dichotomy fails
to grasp the essential plasticity of the human brain –whether of a H. neanderthalensis or H. sapiens.
When viewed at a coarse and long‐term resolution,
there are three features of the archaeological recordthat have led some archaeologists to invoke cognitive
differences between H. neanderthalensis and H. sapiens.
First, despite more than 350,000 years of existence,the extent of cultural and behavioural innovation of the

Neanderthals was limited. Throughout this period oftime they remained as stone‐age hunter‐gatherers witha limited cultural repertoire. In contrast, within the last
200,000 years that H. sapiens has been in existence, thisspecies has invented/discovered agriculture, createdtowns, civilisations, science and so forth. While suchdevelopments are primarily restricted to the last
11,600 years – during the relatively warm, wet andstable Holocene period – the Neanderthals also lived
through similar environmental conditions without
making any fundamental changes in their behaviour.
Anatomical evidence suggests that Neanderthals were
living in conditions of environmental stress; hence
their apparent ‘failure’ to invent bows and arrows, sewn
clothing, substantial shelters appears surprising if they
had the same cognitive capabilities as H. sapiens who
did indeed invent these as means of adaptation tochanging environmental conditions.
Second, there is limited, if any, unambiguous evidence
for the use of visual symbols by Neanderthals. Claims
have been made and debated regarding that coming from
Neanderthal burials, and indeed that burial itself might bea symbolic act. Who manufactured the Châtelperronian‘art’ objects has been a particularly vociferous, butultimately unresolved, debate while the most recent
claims concern the use of pigments by Neanderthals.
The dilemma such claims provides is that it is difficult
to explain the scarcity and ambiguity of such evidence
on the basis of archaeological preservation alone.
Large segments of the Neanderthal archaeologicalrecord are indeed better preserved than that of H.
sapiens, but within the latter evidence for symbolic
behaviour are forthcoming. It is evident that symbolic
communication plays a key role in the adaptation of H.

Neanderthals had a domain‐specific mentality while
H. sapiens possessed the capability for cognitive fluidity,
the latter largely deriving from the use of spoken
compositional language. Wynn and Coolidge later
argued that the Neanderthals had a limited degreeof working memory that constrained their ability atmaintaining multiple ideas in their mind at the sametime – which is in effect another description of anabsence of cognitive fluidity.
While Wynn and Coolidge remained unspecificabout Neanderthal language capabilities, my own view
was that compositional language was absent. How
could it be otherwise when spoken language is the
motor for cultural innovation which is so evidently
absent among the Neanderthals? That left unresolved,
however, how Neanderthals communicated especiallyin light of their vocal tracts and large brains. To address
this my 2005 book, The Singing Neanderthals, proposedthat a sophisticated communication system waspresent that had a greater reliance in variations in tone,
rhythm and pitch and a less reliance on words andgrammar than does compositional language used by H.
sapiens today (and presumably in the past).
The extent to which the domain‐specific model for
Neanderthal cognition remains viable in light of therecently claimed evidence for Neanderthals symbolism
in the form of use of pigments will be considered.
It will be argued that this and other models of
Neanderthal cognition require a greater emphasis on
neuronal plasticity. Indeed, the current archaeologicaldebate between either cognitive differences between H.
neanderthalensis and H. sapiens or differences based on
population and society will be shown to be simplistic
and indeed quite false. One of the key defining featuresof human cognition is the plasticity of the brain that
occurs not only in childhood but also in adults: one
simply can never exclude a consideration of cognition
when trying to explain differences in behaviour
whether within a species or between species. In thisregard we must become more adept at discussing
cognitive variability within both H. sapiens and H.
neanderthalensis, which may enable some degrees ofsymbolic activity to be accommodated in the latter
while maintaining significant cognitive differences
between these species.

The Opening Keynote Lectures

sapiens to challenging environments. Hence in light ofthe environmental stress that the Neanderthals appear tohave experienced it is again surprising that they did notuse symbolic communication more prolifically thanappears to have been the case if they had the capacity
for symbolic thought.
A third reason to argue for significant cognitive
differences between H. sapiens and H. neanderthalensis
is simply the fact that H.sapiens is extant andthe Neanderthals are extinct – and not only the
Neanderthals but all other members of the Homogenus that ever existed. The dispersal of H. sapiens
from Africa at c. 50,000 years ago and their rapid
colonisation of the Old and New Worlds is such aremarkable event that some overriding explanationis required: what could be so fundamentally different
about this particular member of Homo that led to sucha radically different pattern of behaviour to that of all
its relatives and ancestors? Few, if any, suggestions havebeen forthcoming other than an enhanced cognitive
capability, perhaps situated in language.
The counter evidence for cognitive differences
between H. neanderthalensis and H. sapiens rests not
only on the possible existence of symbolic behaviour
by Neanderthals but on three further aspects of thearchaeological and fossil record. First, the large brains
and evolved vocal tracts of Neanderthals appear nodifferent, in essence, to those of H. sapiens. Why would
these costly adaptations exist if they were not delivering
the same degree of advanced cognition? Second,
despite a low level of cultural innovation, the stone tooltechnology of H. neanderthalensis was extraordinarily
sophisticated. Third, H. neanderthalensis endured foran extensive period of time through a wide variety ofchallenging environmental conditions. Could they
have possibly done so without the levels of planningand decision making as found among H. sapiens?
There are, therefore, contradictory strands ofarchaeological evidence regarding Neanderthal
cognition and how, if at all, it differed to thatof H. sapiens. It was an attempt to resolve thosecontradictions and bring archaeological arguments
into line with those in neuroscience and psychologythat I introduced notions of mental modularity. In my
1996 book The Prehistory of the Mind, I argued that029

The Reconstitution of the Neanderthal Brain

Christoph P. E. Zollikofer and Marcia Ponce de León
Anthropological Institute and Museum, University of Zurich, Switzerland

The Opening Keynote Lectures

030

The brain is arguably one of the most complex
organs that evolution has produced. Moreover, it isevident that our own brains stand out from those of
our closest living relatives – the great apes – not onlyby their enormous size, but by their ability to produce
an almost unlimited diversity and complexity ofcognitive, linguistic, and technocultural behaviors. Yet
the exceptionally large brains of our own species Homosapiens are not unique in evolution. Our evolutionary
cousins, the Neanderthals (Homo neanderthalensis), hadbrains that rivaled ours in size. Most notably, it appears
that the large brains of Neanderthals and humans
evolved along independent paths, at least for several
100,000 years.
Which evolutionary and developmental processes,
and which selective and structural constraints, led
to the evolution of these two “brainy” hominin
sister species? Which brain functions did we andthe Neanderthals collectively inherit from our lastcommon ancestor? And how did Neanderthal brainsdiffer from ours in development, structure, function,and cognition? To tackle these questions, we have tofollow various lines of evidence, and apply a diversity of
research strategies. – In this talk, we give an overview
of the current state of Neanderthal brain research, andhighlight major promises and limitations for future
research in paleoneurology.
One major difficulty of paleoanthropology, andespecially of paleoneurology, consists in the fact thatmost fossil specimens are incomplete, fragmented anddistorted, and at least partially embedded in sediment.
Before any biologically relevant analysis can start, fossil
specimens must be reassembled from their isolatedparts; taphonomic distortions have to be corrected, andmissing anatomical regions have to be completed withthe ultimate goal to infer the individual’s morphology
at the time of its death. Over the past decades, we
combined biomedical imaging of Neanderthal
fossils with sophisticated computer graphics tools toreconstruct a large series of fragmentary Neanderthal
specimens, which document the development of
this species from birth to adulthood (Figure 1). The
endocranial cavities contained within these skulls – theso-called endocasts – now represent the material basis
of Neanderthal paleoneurology (Figure 2).
Reconstructing Neanderthal endocasts marks
the first step on the long way toward Neanderthal
brains. Since brains do not fossilize, theirmorphology, and possibly their function, have to
be inferred from the morphology of the endocast.
The morphology of endocasts is highly instructive,
as it provides information about brain size, shapeand structure, which is relevant to infer patterns ofbrain development, structure and function. However,endocranial morphology does not immediately reflect
brain morphology. For example, the form of thebraincase is not only modulated by brain morphology,
but also by the size, shape and orientation of the facerelative to the braincase. Furthermore, the endocast
surface does not provide an immediate picture of thebrain surface, because the space between bone andbrain is filled with meningeal tissue, cerebrospinal fluid
and a wide variety of blood vessels and sinuses. These
structures tend to interfere with, or blur potential
imprints of the brain on the endocast.
To generate reliable estimates of the brain surface of

life Neanderthal brains grew substantially faster thanthose of modern humans. Interestingly, however,
Neanderthals did not reach adult brain size earlier
than modern humans, but on average, they had larger
brains at adulthood. These differences have various
implications for the life history of Neanderthals. At
first sight, it appears that their fast-growing brains fit
into the widely held picture that Neanderthals lived
according to a “live fast – die young” scheme. However,from a wider comparative perspective, large adultbrains imply a longer life and even slower reproduction
than in modern humans.
The marked differences between Neanderthal
and modern human endocranial shape indicate that
there have been major species-specific differencesin brain development before birth. Following thedevelopmental trajectories of Neanderthal and modern
human endocasts after birth indeed shows that theseprenatal differences are carried over into differencesfrom infancy through to adulthood. As mentioned,
the relationship between the endocast and the brain iscomplex, such that any inferences on the structural andfunctional development of the brain have to be takenwith great care. During the early 19th century, it wascommon practice among the “phrenologists” to inferpersonality traits from the individual shape of a person’s
braincase. While modern neurobiology has debunked
this simplistic path of inference, paleoneurologists
are still tempted to establish direct links between theendocast and the brain, inferring major differences in“higher” cognitive functions between Neanderthals
and ourselves.
What, if anything, can then be said about
Neanderthal brain structures, brain development,
and ultimately about brain function and cognition?
One possibility is to infer differences in cognition
and behavior between Neanderthals and modern
humans from the archaeological record, which, tosome extent, reflects the “behavioral output” of theirrespective brains. However, it is difficult to tell apartintraspecific technocultural differences from speciesspecific differences. Another possibility is to widen thescope of paleoneurology, including living great apes ascomparative species, and adopting the methodologicaltoolkit of modern neurobiology. Magnetic Resonance

The Opening Keynote Lectures

a Neanderthal from its preserved endocranial cavity,
comparative data from living species are needed.
Biomedical imaging methods such as CT and MRI
now permit the acquisition of bone and brain datafrom the same subject, not only in humans but
also in our closest living relatives, the chimpanzees
and other great apes. We developed an array of
endocast-brain mapping procedures, which combine
computer-assisted fossil reconstruction with soft
tissue reconstruction methods used in forensics. Thisapproach now permit inferences on the size, shape andsurface structure of Neanderthal brains.
Only very few Neanderthal individuals are available
to document the decisive early phases of brain
ontogeny, where slight differences in rates of growthand development produce significant differences inbrain size at adulthood. The decisive information about
these phases comes from the newborn Neanderthal
from Mezmaiskaya Cave (Russia) and from the two 1.0
and 1.5 year-old children from Dederiyeh Cave (Syria)
that have been excavated by a Japanese-Syrian teamled by Professor Takeru Akazawa.
Our analyses show that the brains of newborn
Neanderthals and humans were similar in similar size(400 ccm), but clearly different in shape. Similarity
in size is likely to reflect similarity of developmental
constraints in both species. One such constraint
results from the fact that, during birth, the head of
the fetus has to pass through the constricted pelvic
outlet of its mother. In modern humans, neonate
head dimensions closely match the dimensions of
the maternal pelvis, which is a major cause of birthcomplications. Our computer-assisted reconstructionof the only fairly well-preserved female Neanderthal
pelvis (the specimen from Tabun, Israel) indicates that
Neanderthal mothers had a more spacious pelvis thanmodern human mothers. However, it appears thatbirth in Neanderthals was as complicated as in modern
humans, because Neanderthal newborns had evenlarger heads than modern human newborns. This was aconsequence of the peculiar shape of the Neanderthal
braincase, and their comparatively large face already atbirth.
Tracking brain growth trajectories from birth toadulthood shows that during the first two years of031

Imaging (MRI) now permits non-invasive acquisitionof structural and functional brain data from humanvolunteers and anesthetized great apes. Furthermore,
recent methods of genomics and transcriptomics
permit investigation of the genetic networks thatmediate brain growth and development. Thesemethods have already yielded surprising insights into
differences and commonalities between human andgreat ape brain development, structure and function,and into the genetic underpinnings of these differences.
Establishing structure-function relationships between
the genome and the phenome is a dauntingly complex
endeavor, especially if it is about brains. However,given our steadily increasing knowledge about the
Neanderthal genome and phenome, it is a research
path worth to be followed. Wherever Neanderthal
paleoneurology will bring us in the near future, wehave to keep in mind that this endeavor will influence
our views on Neanderthal brains, and vice versa.
Reconstituting the Neanderthal brain will change theway in which we think about the Neanderthals andabout ourselves.

The Opening Keynote Lectures

032

Figure 2. Endocasts of a Neanderthals (left) andmodern humans (right). Top: the 4 year-old Neanderthal
child from Devil’s Tower, Gibraltar (left) and a modern
human child with similar age (right; clinical data). Bottom:
the endocast of the Le Moustier adolescent Neanderthal(left) and that of an adult modern human (right).

Figure 1. Reconstructing the early development of
Neanderthals. (left: the Mezmaiskaya neonate; right: theDederiyeh 1 child, with an estimated age at death of 1.6
years).

Or

Oral Presentationsa

The Process of RNMH: Global and Comprehensive Explanatory Models
Learning Behaviors in Prehistoric and Modern Humans
Human-specific Learning Strategies and Cultural Evolution
Towards Digital Reconstruction of Fossil Crania and their Brain Morphology
Neuroimaging Approach to the Learning Hypothesisl

RNMH2012

The Replacement of Neanderthals by Modern Humans
in EuropeJoão Zilhão

Seminari d'Estudis i Recerques Prehistòriques, University of Barcelona, Spain

The Process of RNMH: Global and Comprehensive Explanatory Models

034Over the last two decades of the 20th century,
debates on the emergence of European modern
h u m a n s a n d t h e f a t e o f t h e Ne a n d e r t a l stended to revolve around the polar alternativesof “Multiregionalism” (in its original, strict
formulation, implying that European moderns
had evolved from local, ancestral populations of
Neandertals) and “Recent African Origin” (inits original, strict formulation, defining modern
humans as a new species with a birthplace in
East Africa some 150,000 years ago, one that
subsequently went on to colonize the rest of
the world through a process of Out-of-Africa
expansion in the course of which Eurasian archaic
humans, namely the Neandertals, would have gone
extinct without descent).
A minority position, the “Assimilation” model,
accepted recent Out-of-Africa migration and/or
genetic diffusion but viewed Neandertals not asa different species but rather as a geographicallyisolated Eurasian variant of Homo sapiensthat ended up being assimilated in the wider
human gene pool after 40,000 years ago as aresult of demographic and/or natural selection
processes operating in a context of significant
population admixture. Paleontological, geneticaland archeological research has shown such
“Assimilation” views to provide the closest match
to the revised and/or augmented empirical baseconcerning the replacement of Neandertals bymodern humans gradually accumulated over thelast 15 years.
Direct dating of the fossils that were thought torepresent the earliest European modern humans— and, through their lack of anatomicallyarchaic features, supported replacement-withno-admixture of the aboriginal Neandertals —showed that they were all significantly younger,
some times of recent Holocene age. Conversely,
the newly discovered or restudied fossils dated
to within five to ten millennia after the time ofcontact (e.g., the Lagar Velho child, the Oase 1
mandible, and the Oase 2 cranium) were all shown
to present anatomically archaic, if not Neandertaldiagnostic features. These morphological mosaics
were interpreted as indicating admixture at thetime of contact and, despite much controversy, theNeandertal genome project eventually produced
corroborating evidence; namely, that 1-4% of
the genome of extant Eurasians is of Neandertal
origin.
At the same time, archeological research
provided evidence that, in the behavioral realm,
late Neandertals had been as “modern” as their
African contemporaries, thereby removing putative
cultural barriers to interbreeding from the range of
mechanisms that conceivably could have prevented
admixture from occurring, while showing thatsome of the Upper Paleolithic innovations that
“Recent African Origin” views interpreted asbrought in by immigrating modern humans shouldin fact be credited to the Neandertals (namely, theuse of pendants made of ivory, bone or tooth, aswell as, possibly, the earliest, abstract/geometriccave paintings).

Cultural and Biological Transformations in the Middle
Pleistocene Levant: A View from Qesem Cave, Israel

Ran Barkai and Avi Gopher

Department of Archaeology, Tel Aviv University, Israel

H. erectus (senso lato) was present in the Levant
during a period of over one million years, between
1.4 my and 400,000 years ago, and created theAcheulian cultural complex. We suggest thatduring this long period of time H. erectus wasdependent on elephants (and especially their fat)for his survival.
The disappearance of elephants from the Levantsome 400,000 years ago coincides with the end of
the Acheulian complex and the appearance of a
new and innovative local cultural complex – theLevantine Acheulo-Yabrudian. As is evident fromthe dental remains recently found in the AcheuloYabrudian site of Qesem Cave, the hominins
occupying this cave were not H. erectus but wererather most similar to later Modern populationsin the region (e.g., Skhul/Qafzeh). This hominin
exhibits a new set of behavioral and cultural
traits (e.g., habitual use of fire; meat sharing;
innovative cutting tools; flint quarrying; lithic
recycling etc.). Archaeological evidence worldwide seems to associate H. erectus with large andmediumsized game, most conspicuously elephants
whose remains are commonly found at Acheuliansites throughout Africa, Asia, and Europe. In
some instances elephant bones and tusks were alsotransformed into shaped tools, specifically artifacts
reminiscent of the characteristic Acheulian stonehandaxes. In Africa, H. sapiens appears around 200
kyr ago, most probably replacing H. erectus and/or H. heidelbergensis. Early African H. sapiens sites
are devoid of elephants. The presence of elephants
in many African Acheulian sites and their absence

The Process of RNMH: Global and Comprehensive Explanatory Models

In this presentation we provide our interpretation
for the circumstances leading towards theevolutionary replacement of Homo erectus (sensolato) by a new hominin lineage in the Middle
Pleistocene Levant, some 400,000 years ago.
In short, our hypothesis is that dietary stresscaused by the disappearance of elephants triggered
the extinction of H. erectus, a hominin species
highly dependent of the consumption of calories
from very large animals, and the evolution of a
new hominin lineage that was better adapted to
the new diet constraints of hunting larger numbersof smaller and faster animals in order to provide
sufficient caloric intake to compensate for theloss of the elephants. The biological replacement
took place in tandem with significant culturalchanges and the appearance of a new, unique andinnovative cultural complex in the Levant.
It is our contention that the appearance of a newand creative set of behaviors 400,000 years ago in
the Levant must have been supported by innovative
cultural transmission mechanisms, very different
than the ones practiced during earlier Lower
Paleolithic times. Advanced learning behaviors andabilities must have played a significant role in the
adoption and assimilation of new hunting, meat
sharing, flint procurement and flint production
strategies, as well as in the earliest habitually use offire. These new cultural traits characterized humanexistence in the Levant for a long period of over
200,000 years, to be later replaced again by the
Middle Paleolithic cultural complex created byboth Modern humans and Neanderthals.

035

The Process of RNMH: Global and Comprehensive Explanatory Models

036

from later Middle Stone Age sites evoked an
overkill hypothesis, which was never convincingly
demonstrated. Thus no link was proposed, in thecase of Africa, between human evolution and theexclusion of elephants from the human diet, andno evolutionary reasoning was offered for the
emergence of H. sapiens in Africa. We suggest thatthe disappearance of the elephants, created a needto hunt an increased number of smaller and fasteranimals while maintaining an adequate fat content
in the diet, and this was the evolutionary drive
behind the emergence of the lighter, more agile,and cognitively capable modern hominins in theLevant.
Qesem Cave is a Middle Pleistocene site in
Israel, dated to 420,000-200,000 years ago (Barkai
et al., 2003; Gopher et al., 2010) and assigned
to the the Acheulo-Yabrudian cultural complex(AYCC) of the Lower Palaeolithic Levant. The
cave reveals a rich and well-preserved array oflithic and faunal remains as well as human teeth. It
provides a good context in which to test hypotheses
concerning the intriguing liaison between theenvironment, culture, and biology. Here wesummarize a decade of research suggesting thatthe unique cultural transformation seen at Qesem
Cave might actually be related to local human
evolutionary processes and the appearance of a
new hominin lineage.
As attested at Qesem Cave, the AYCC is aunique, local cultural entity clearly differing fromthe preceding Acheulian and the proceeding
Mousterian. It shows a suite of innovative
behaviors including: the habitual use of fire;
hearth-centered activities and functionally distinct
activity areas; sophisticated raw material acquisition(flint quarrying); intensive and systematic blade
production employing an efficient, innovative, andthoughtful technology; a noticeable presence of'ahead of their time' Quina scrapers; and intensive
flint recycling activities.
The 9.5 m stratigraphy of Qesem Cave offers adetailed background of sediment deposition that
occurred as the karstic chamber cave was aging.
Various environmental proxies correlate with

global climatic timelines (stages 11-7) and more
specific data reveal the dynamics of a changing
ecology within generally Mediterranean regimes offorest vegetation zones.
Cave inhabitants hunted cooperatively, bringing
body-parts of fallow deer back to the cave, whichwere then butchered, shared, and – as evidenced
by the use of fire throughout the cave’s 9.5 m
deep stratigraphy and the many burnt bones –eventually barbecued (Karakanas et al., 2007;
Stiner et al. , 2009, 2011). Abundant cutting
tools were systematically produced at the site,
most significantly flint blade knives made by an
innovative and thoughtful technology (Barkai
et al., 2005, 2009; Shimelmitz et al., 2011).
Our lithic analyses and the study of use-wear
signs on flint artifacts indicate a set of cutlery
manufactured to handle the different stages ofbutchering, defleshing, and meat cutting (Barkai
et al. ,2010; Lemorini et al. 2006). A study ofdental remains (Hershkovitz et al., 2011) indicatedthat the hominins inhabiting Qesem cave were
not H. erectus but rather most similar to later
modern populations (e.g., Skhul/Qafzeh) of
this region, notwithstanding some Neanderthal
affinities as well. Based on a bio-energetic model
it was recently suggested that the biological andcultural developments attested at Qesem Caveoccured in the Levant some 400,000 years ago as
an adaptation to new dietary constrains caused
by a faunal turnover that led to the exclusion ofelephants from the human diet (Ben-Dor et al.,2011). These results are consistent with the recently
published innovative model for the evolution of
the human Pleistocene populations of Europe(Bermúdez de Castro and Martinón-Torres 2012),
suggesting the Levant as the Central Area of
Dispersals of Eurasia (CADE), an “origin region”
for human species biodiversity. Acknowledging weare challenging a well-established paradigm, our(as yet) insufficiently comprehensive hypothesis isoffered with the intention of opening new vistas
for discussion.

Middle Pleistocene Qesem cave, Israel," Quaternary
Geochronology 5: 644-656.
Hershkovitz, I., Smith, P., Sarig, R., Q uam, R.,
Rodríguez, L., García, R., Arsuaga, J-L., Barkai, .R.,
and Gopher, A. 2011. "Middle Pleistocene dental
remains from Qesem Cave, Israel," American Journalof Physical Anthropology 144: 575-592.
Karkanas, P., Shahack-Gross, R., Ayalon, A., BarMatthews, M., Barkai, R., Frumkin, A., Gopher,A., and Stiner, M.C. 2007. "Evidence for habitualuse of fire at the end of the Lower Paleolithic: Site
formation processes at Qesem Cave, Israel," Journalof Human Evolution 53: 197-212.
Lemorini, C., Gopher, A, Shimelmitz, R., Stiner, M., andBarkai, R. 2006. "Use-wear analysis of an Amudian
laminar assemblage from Acheuleo-Yabrudian
Qesem Cave, Israel," Journal of Archaeological Science
33: 921-934.
Shimelmitz, R., Gopher, A., and Barkai, R. 2011.
"Systematic blade production at Late Lower
Paleolithic (400-200 kyr) Qesem Cave, Israel,"
Journal of Human Evolution 61: 458-479.
S t i n e r, M . , G o ph e r, A . , a n d B a rk a i , R . 2 0 0 9 .
"Cooperative hunting and meat sharing 400-200 kya
at Qesem Cave, Israel," Proceedings of the National
Academy of Sciences 106: 13207-13212.
Stiner, M., Gopher, A., and Barkai, R. 2011. "Hearthside socioeconomics, hunting and paleoecology
during the late Lower Paleolithic at Qesem Cave,
Israel," Journal of Human Evolution 60: 213-233.

Figure 2. Excavations at Qesem
Cave during the 2011 fieldseason.

Figure 1. Location map of
Qesem Cave.

Figure 4. Butchering experiment using
replicas of recycled small flakes from
Qesem Cave.

Figure 3. Flint blades from
Qesem Cave.

The Process of RNMH: Global and Comprehensive Explanatory Models

Bibliography
Barkai, R., Gopher, A., Lauritzen, S.E., and Frumkin,A. 2003. "Uranium series dates from Qesem Cave,
Israel, and the end of the Lower Palaeolithic,"
Nature 423: 977-979.
Barkai, R., Gopher, A., and Shimelmitz, R. 2005.
"Middle Pleistocene Blade Production in theLevant: An Amudian Assemblage from Qesem
Cave, Israel," Eurasian Prehistory 3: 39-74.
Barkai, R., Lemorini, C., Shimelmitz, R., Lev, Z., Stiner,M., and Gopher, A.. 2009. "A blade for all seasons?
Making and using Amudian blades at Qesem Cave,
Israel," Human Evolution 24: 57-75.
Barkai, R., Lemorini, C., and Gopher, A. 2010.
"Palaeolithic cutlery 400 000–200 000 years ago:
Tiny meat-cutting tools from Qesem Cave, Israel,"
Antiquity 84: 325.
Ben-Dor, M., Gopher, A., Hershkovitz, I., and Barkai,
R. 2011. "Man the fat hunter: The demise of Homoerectus and the emergence of a new hominin lineage
in the Middle Pleistocene (ca. 400 kyr) Levant,"
PLoS One 6(12):e28689, doi:10.1371/journal.
pone.0028689.
Bermúdes de Castro J-L & Martinón-Torres, M. 2012.
A new model for the evolution of the human
Pleistocene populations of Europe. QuaternaryInternation.
Gopehr, A., Ayalon, A., Bar-Matthews, M. Barkai, R.,
Frumkin, A., Karkanas, P., and Shahack-Gross, R.
2010. "The chronology of the Late Lower Paleolithic
in the Levant: U series dates of speleothems from037

From the Beginning: Tracing the Patterns of Cultural
Changes of Early Homo sapiens in Africa, West Asia,
and Europe
Seiji Kadowaki1 and Katsuhiro Sano2

1Nagoya University Museum, Nagoya University, Japan 2Department of Archaeology, Tohoku University, JapanThe Process of RNMH: Global and Comprehensive Explanatory Models

038This paper derives from our archaeologicalinvestigations on prehistoric learning behaviors,
particularly those involved in anthropologicalprocesses of the replacement of Neanderthals bymodern humans (hereafter RNMH). As part ofthe studies conducted by Research Team A01
of the RNMH project, we have been compilingarchaeological data relevant to the RNMH process
under two purposes. The first is to collect andorganize updated material evidence regarding
the RNMH in a comprehensive manner, and thesecond is to obtain insights into prehistoric learning
behaviors through the observation of diachronic andgeographic patterns of cultural variability. Using partof the archaeological database compiled since 2010,

i.e., the beginning of the RNMH project, this paper
examines chronological and geographical culturalvariability during the Middle and Later Stone Age(hereafter MSA and LSA respectively) in Africa
and the Middle and Upper Palaeolithic (hereafter
MP and UP respectively) in west Asia and Europe,and then discusses some implications of thisarchaeological evidence on the learning behaviors by
early Homo sapiens.
The temporal and spatial range covered by the
database of this project is set from ca. 300 to 20 kya
in Africa and Eurasia, which is broader than those
directly related to the RNMH events per se. Thisis because our aim is not just to trace the timings
and frontiers of the RNMH events on the a priori

Figure 1. Cultural chronology of the MSA and LSA in Africa and the MP and UP in West Asia and Europe. Colors ofcultural units indicate general technological characteristics (Clark's technological mode) of constituent lithic assemblages.

Figure 1 (continued).

Neanderthals and modern humans. Although
theoretical links between learning strategies andpatterns of cultural changes are proposed on thebasis of cultural evolutionary theories, an attempt
for their empirical testing entails a number ofchallenges. First of all, archaeological remains thatcan be securely associated with hominin taxa arelimited. Moreover, it is difficult to reliably assess the
speed and cumulativeness of cultural changes, which
are considered significant aspects in the cultural
evolutionary theory studied by Research TeamB01 of the RNMH project, because of numerous
chronological issues and variable definitions ofprehistoric cultures. Because these challenges
cannot be readily resolved, the present study would
rather address and clarify these problems throughempirical examinations of chronological andgeographic distributions of archaeological cultures.
We will then propose parsimonious interpretationsof the patterns of cultural changes during the MSA
and LSA in Africa and the MP and UP in west
Asia and Europe in terms of prehistoric learning
behaviors by early Homo sapiens and Neanderthals
to contribute to the RNMH project.
The Process of RNMH: Global and Comprehensive Explanatory Models

assumption that such anthropological events are
directly reflected in archaeological records, but toorganize archaeological data available from thetime periods and geographic areas, where modern
humans presumably emerged and dispersed withprobable replacement of preceding populations,
including Neanderthals. Because much remains to
be clarified in the association between archaeologicalremains and hominin taxa, we put more emphasisin this study on the systematic presentation andassessment of archaeological data broadly related
to the RNMH process rather than attempting
to provide definitive archaeological answers toanthropological problems of the RNMH.
On the other hand, the broad temporal andspatial ranges of our archaeological investigations
are also expected to provide us with sufficient sizeof data to examine diachronic and geographic
cultural variations by modern humans and otherpreceding hominins, particularly Neanderthals.
This analysis is intended to help us draw some
implications on prehistoric learning abilities/
behaviors, which constitute the main research
theme of the RNMH project that hypothesizes thedifferences in learning abilities/behaviors between

039

Thinking the Beginning of Early Upper Paleolithic in
Siberia and Central Asia

Hirofumi Kato and Masaki Naganuma
Center for Ainu & Indigenous Studies, Hokkaido University, JapanThe Process of RNMH: Global and Comprehensive Explanatory Models

040

The aim of this report is to provide an overview
of the Early Upper Palaeolithic cultural complexes
in the Northern Eurasia region where Neanderthaland Homo sapiens both had invaded from outside
areas and achieved development and evolution of
their cultures.
We will examine the background of the Early
Upper Palaeolithic cultural formation processes
based on paleo-environmental conditions andtheir cultural characteristics. As next step, we
compare and consider formation processes of theUpper Paleolithic by tracing the aspects of Midleto Upper Palaeolithic transitions.Especially, we
focus on the following 4 regions where provide
some important achievements nowadays; Ural
Mountains, Altai mountains, Central Asia andthe Baikal Siberia. Using as basic data fromour "NeanderDB" still in progress building by
us, this study investigates some hypotheses anddemonstrates notably viewpoints and additional
new discoveries and other information.
Aspects of our basic materials are as follows:(1) At present, the paleoanthropological materials
have concentrated on the Altai mountains
region. The human fossils from the Denisova andOkladnikov caves suggestthat there were threehuman populations in the MP-UP transitonal
period; Homo sapiens, the Neanderthals, andthe Denisovians, paleoanthropologcaly andmeroculargeneticaly. While in the Central Asia,
some paleoanthropological materials were also
found and they have certain importance to tracesof the Neanderthal's migration to the Southern

Siberia, and they also had effects on the molecular
genetic data.
(2) The archaeological materials suggest there p e a t e d i nv a s i on f rom We s t : t h e Pe b b l etool cultures, Keilmessar groups, and Eastern
Gravettian in the Ural Moutains. On the other
hand, at the Altai mountains, after the invasion
by Mousterian people during around 300 ka, thehypothesis of local technological developmentof litihic industries and gradual transition from
Middle to Upper Palaeolithic have been insisted.
In the Baikal Siberia (both of Cis and Trans
Baikal) region, it seems that there were transitions
from non-Mousterian local Middle Palaeolithic
to Upper Palaeolithic, but its ancestries are still
unknown. For considering distribution andancestries of these Middle Palaeolithic cultures,
it is notable to compare with Middle Palaeolithic
aspects in the Central Asia, especially theirchronological contexts as well as relationshipbetween anthropological data and several variantsof the Mousterian.
(3) As for the leaning behaviors, it had been
pointed out that the elements of behavioral
modernity such as potable arts (venus, figurines ofanimals), personal ornaments, and ochers, emergedin Northern Asia older than European regions. In
our opinion, it will be important that to compare
difference of site locations, hunting games,
exploitation areas before and after the emergenceof elements showing behavioral modernity. It
would be necessary to create exceeding direct
obser vation for the background of making

animal designs. In addition, personal ornaments
might be connected with the emergence of more
complicated innovations on the sharing othergroups and reciprocal alliance.
(4) As for the emergence of religions, the new
discoveries suggest the burial contexts of Venus
and animal figures; from Zaraisk and Mal’ta Upper
Palaeolithic sites. If classical Upper Palaeolithic
after 20 ka is defined by appearance of developing
movement inner world as well as just living in
actual world, the Early Upper Palaeolithic would

be its infancy. The factors of the formation of
spiritual cultures might suggest that unstabletransitional period had drastically changed
environment, culture and human population in
Northern Eurasia.

The Process of RNMH: Global and Comprehensive Explanatory Models

Figure 1. Distribution of the MP, MP-UP transitional, and EUP sites in Siberia and Central Asia.

041

Investigation of Past Climate and Vegetation for
Understanding the Replacement of Neanderthals byModern Humans
Ayako Abe-Ouchi1, Yusuke Yokoyama1, Hodaka Kawahata1,
Takashi Oguchi2, and Minoru Yoneda3

1Atmosphere and Ocean Research Institute, The University of Tokyo 2Center for Spatial Information Science, The University of Tokyo
3The University Museum, The University of Tokyo

The Process of RNMH: Global and Comprehensive Explanatory Models

042

The impact of climate change for the extinction of
Neanderthal and other archaic human species related
the dispersal of modern human (Homo sapiens) is
intriguing but still challenging topic for archaeologists
and anthropologists. There are several reasons why
this is difficult to compare the climatic data fromdifferent kinds of media including ice cores, lake andocean sediments and stalagmites, and the distributionof human species. In this project, we would like tocontribute to provide some basic information for the
temporal and spatial variability of climatic zones andancient human species which could be recognized bydifferent stone industries.
The temporal change of climate change wasreconstructed from oxygen isotopic ratios inforaminifera from the benthic sediment cores, andoxygen isotope and other climatic proxies recordedin ice sheets from Greenland and Antarctic. That
showed very drastic cycle of cooling and warming
events in the late Pleistocene in which periods themodern human dispersed from Africa to Eurasia.
However, the direct comparison between the eventsin human evolution and climatic events had beendifficult because of dating methodology. The ice coredata which are accepted as proxy of global climatechange provide us more detailed temporal changesuch as Dansgaad-Oeschger events and Heinrich
events clearly. The timing of each event are generally
determined by counting annual rings recorded in ice
core.
On the other hand, most archaeological dates
regarding to the extinction of Neanderthal were
determined by radiocarbon dating which is basedon the reduction of radiocarbon from the "100%modern carbon" which fluctuate by change of theintensity of cosmic ray and geomagnetic field. Until
recently, the fluctuation of radiocarbon activity inthe late Pleistocene could not be estimated and theradiocarbon dates in this period theoretically containseveral hundreds or thousands years uncertainties.
By combining a series of data from marine cores andcorals, the calibration curve for radiocarbon dates was
expanded to 50 ka by Intcal09 dataset. This means we
can compare the timing of events in archaeologicalrecords can be investigated in the history of global
climate change at last. We are building a databaseof radiometric dates regarding to the extinction of
Neanderthal and dispersals of modern humans inlight of methodological evaluations. Some of key
data must be taken away because of insufficient
preparation and/or measurement. The selected
data will give us more concrete and reliable map of
distribution of different human species in time andspace.
Second point which we have to overcome is thespatial variability of climatic and environmental
conditions. As mentioned above, many geochemical
proxies have showed the regional and global historyof climate change in detail but the climatic map inthat period is not easy to obtain for archaeologicaland anthropological discussions. For our project, oneof authors heading a team of paleoclimatologists
and geophysicists to make a series of climatic
distribution maps to compare with the distributionof different human species. A coupled atmosphereocean general circulation model, named the Model

The Process of RNMH: Global and Comprehensive Explanatory Models

for Interdisciplinary Research on Climate (MIROC)
was applied, which have produced results for the
distant past including 6 ka, LGM of 21 ka, and midPliocene 3 Ma. Some additional models for thegrowth of ice sheets and the dynamics of surface
vegetation were also included.
Finally, we compare the results from the human
distribution based on radiometric dating database
and the simulated pale climatic maps in order tounderstand the climatic impact on the replacementof Neanderthal and other archaic humans by our
species. For this purpose, we improve a ecologicalprogram for niche modeling. For example, we apply
the MaxEnt program using maximum entropy model
to see the suitable conditions including climate,geography and biozone for different human species.
For applying this model, we use improved higher
resolution (e.g., 60 km mesh) map for paleoclimate
and the distribution of probability of calibrated
radiocarbon dates. The preliminary results shows
overlap and some differences in possible ecologicalniches between Neanderthal and modern human. We
will investigate the temporal change of their suitableconditions in light of change in cognitive ability and/or way of life.

043

Cultural Transmission, Institutional Continuity and
the Persistence of the Mousterian

Steven L. Kuhn

School of Anthropology, University of Arizona, Tucson, USA

The Process of RNMH: Global and Comprehensive Explanatory Models

044

By all accounts Mousterian or EurasianMiddle Paleolithic material culture represents an
extremely successful set of hominin adaptations.
Mousterian technologies were used for more than
200,000 years by groups living in diverse habitats
through several glacial/interglacial cycles. Yet theyalso disappeared surprisingly rapidly following
the expansion of anatomically modern Homosapiens into Eurasia. In fact, the same factorsthat underpinned the apparent success of theMousterian technologies and culture may have
hastened their eventual disappearance. Genetic
evidence indicates that Middle Paleolithic
populations, though widespread, were small andfragmented into numerous local demes. Middle
Paleolithic foragers fed at a high trophic level,
which would have facilitated expansion into
empty habitats but also would have kept absolutedensities low. These demographic conditions help
explain the apparent continuity in Mousterianculture across great expanses of space and time.

Small, fragmented populations limit the rates atwhich innovations appear and spread and constrain
the scale of random drift from ancestral states. At
the same time, unstable populations inhibit thedevelopment of robust social networks and othercultural institutions that can store latent cultural
information, a necessary pre‐requisite to cumulative
culture change. Equally important, it is difficultfor dispersed, fragmented populations to resist
invasion. Any cognitive or cultural characteristics
which led to greater continuity in early Homosapiens populations and cultural institutions would
have helped them establish a permanent presencein Eurasia. The presence of invading groups wouldin turn have disrupted the already fragile social
networks of Middle Paleolithic populations,disrupting lineages of cultural inheritance. Due tothe more volatile nature of cultural transmission,
the cultural heritage of indigenous groups was
swamped even more quickly than their genes.

Evolutionary Development of Learning and Teaching
Strategies in Human Societies

Hideaki Terashima
Kobe Gakuin University, Japan

1. The long duration of childhood: Modern
hunter-gatherers have a quite long childhood from
infancy to adolescence and semi-adult in contrast
to the Neanderthals who are said to have had only
a limited period of childhood. During childhood
children do not need to contribute to subsistence
work and they can freely engage in play activities.
Childhood is the time for brain development anddefinitely important period for learning.
2. During infancy, some innate learning processthat responds to teaching behavior of adults
begins, but from some time after weaning, children
rarely receive direct teaching from adults. Theyleave their mothers or caregivers and join intoplay groups consisted of multiage children, where
they learn various knowledge and skills throughplay. Particularly pretended activities such as playhunting are very important to obtain knowledge
and skills not only for subsistence work but also
to obtain an understanding of the nature itself.
Nature affords the environment of learning with its
diversity, complexity and dynamism which arousemuch curiosity and desire for learning in children.
3. Children usually learn more by observing andimitating others on their own during play withother children than by receiving instructions
from others. Learning through participation
into communal activities is almost sufficient toobtain knowledge and skills for daily activities.
Adults usually avoid giving strong instructions tochildren except for some occasions. This style ofautonomous learning seems to foster creativity andadaptability in children.
4. Collage expression, picture drawing and HandTest carried out among the Baka children in
Cameroon suggested their interests, creativity,
attitude toward new experiences, and resilience,that is, the flexible attitude to difficult conditions.
The cognitive flexibility was also suggested inchildren's object making activities. Psychologicalexperiments for young children to investigate theLearning Behaviors in Prehistoric and Modern Humans

As a team of cultural and biologicalanthropologists and developmental psychologists
we aim at understanding the characteristics oflearning behavior and learning strategies of modern
hunter-gatherers through fieldwork to obtain clues
to the replacement of Neanderthals by modern
humans. We pick up the following three points asmain fields of research: 1) hunting and gathering
as a way of life, 2) children and childhood, and3) children's play. These points are chosen basedon the following ideas. First, we need to explore
how people learn in hunting and gathering lifethat was the only lifestyle of humans during the
replacement. Second, childhood is the most critical
period for learning not only to get knowledge andskills for daily life but also to provide foundation
for later development of innovative behavior. Third,
in a society without school education, children
learn mainly through daily activities, particularly
playing in play groups. The following are someresults of the study to the present time.

045

ability to comprehend visual signs and pictures
showed their intuition and representation to signs
and visual reception along with their flexibility.
5. Verbal instructions are not so much used fortransmission of knowledge and skills as for social
norms and values such as sharing, cooperationand friendly behavior. For those invisible things
language is employed in various manners. Many
metaphors are used in stories containing a varietyof anthropomorphic images. Anthropomorphism,
sometimes considered to be a primitive way of
thinking, seems to be related to the developmentof various cognitive abilities and used as a uniqueand excellent method for representing world
reality. Also ritual ceremony such as initiation rite
works as an occasion to teach social values andmorals that are rather difficult to teach only withverbal instructions.

Learning Behaviors in Prehistoric and Modern Humans

046

6. During adolescence daily activity of young
people, males particularly, increases very much inaccordance with growth spurts of the body. They
explore deeply into the environment and expand
the scientific knowledge of nature, in particular,
animals, plants and other things related tosubsistence work and also obtain an understandingof how nature works. Young adults visit distantgroups from time to time, one of the goals being
to find a mate. Such a visit intensifies theirsocial network and brings about an occasion fortransmission of information and innovation.
7. The cultural innovation in hunting andgathering societies covers very wide domains frommaterial culture such as foraging tools, clothing
and housing to non-material items such as ritual,song, dance, play, and body decoration. Mostof those innovations are not made individuallyin each group but transmitted from place toplace. Innovation becomes recognized and fixed
through transportation to other groups where it
may serve to give birth to new ones. In modern
hunter-gatherers innovation is not only related toindividual ability to innovative behavior but also

closely connected to the social learning system.
8. The band, a daily living organization of huntergatherers, is characterized by flexible membershipand fluidity. Bands are connected to each other by
their social network. People frequently pay visits
to relatives and friends through such networks toprovide opportunities for the exchange of people
and goods. Those extensive social networks andfrequent movement of people make the baseof cultural innovation. Comparison between
Neanderthals and modern humans in terms of therange of people's mobility may help to consider the
degree of Neanderthals' cultural innovation.
9. Cognitive sciences have revealed the importanceof metacognition and meta-learning in modern
humans. Our anthropological fieldwork alsosuggests the usefulness of "teaching without
teaching" as a method of education. Metacognition
and meta-learning presuppose higher cognitive
ability and brain function. The large morphologicaldifference of the brain between Neanderthals
and modern humans suggests some importantfunctional differences between them, one of which
may be the difference of learning ability on thelevel of metacognition.
10. The evolution of learning ability depends
not only on a single factor but on the synthetic
development of various factors in the following
three domains: cognitive abilities, knowledge
about the nature, and social system. Investigationof learning behavior in modern hunter-gatherers
so far suggests that factors in those three domains
are interconnected and related to each other tocreate their unique learning strategy. Collaboration
between anthropological fieldwork, developmental
psychology, educational sciences, cognitive sciences
and primatology should be continued to elucidate
the nature of the human learning system.
Here we have a tentative sketch of the evolution of
the human learning capacity:(1) Australopithecus and other ape-men: Thosehuman ancestors were almost at the same level
as contemporary great apes like chimpanzees
and bonobos in terms of cognitive ability andsubsistence skills. During this period they carriedout social learning such as obser vation andimitation (emulation), but with slight culturaltransmission between generations.
(2) Homo erectus and Neanderthals: There weresocial transmission of cultural heritage based on
excellent social learning. In Neanderthals there also
might be teaching behavior in order to reproduce
the elaborate stone tools with almost the same
shape for hundreds of generations. However, in
this stage development of innovative behavior andsocial networks was minimal. Also the cognitive
ability to handle the matter of metacognition
seems to be insufficient.
(3) Homo sapiens: A big development took placein learning behavior due to the development ofcognitive ability, in particular, metacognition. The
knowledge on the nature expanded enormously
and brought about the development of imaginativeand symbolic thoughts. Flexible and creative
individual learning along with rapid and accuratesocial learning produced a powerful learning
strategy in modern humans. The development ofsocial network and mobility of individuals brought
frequent cultural exchange which led to further
creation of new ideas and technology.
Learning Behaviors in Prehistoric and Modern Humans

047

New Insights into the Skill Learning Processes in the Lithic
Production: An Analysis of the Refitted Material from the
Kyushirataki 15 Site in Hokkaido, Northern Japan
Jun Takakura

Hokkaido University, Japan

Learning Behaviors in Prehistoric and Modern Humans

048

Refitted portions of lithic tools and production
debitage found together can be viewed asreflections of the past knapping activities carriedout at a specific time, at the place of discovery. An
analysis on the spatial patterning of the refitted
materials may reveal the past human behavior
involving the skill transmission process. If
numerous pieces of artifacts including cores, bladesor microblades, retouched tools, and debitage
are successfully refitted, we can understand how
the past stoneworkers operated and organized
their actions along with technological choices
and decisions, from the initial acquisition of raw
materials, through tool manufacture, use, anddiscard (Bleed, 2002; Takakura, 2010). The study oflithic refitted materials, especially paying attention
to a difference of the knapper’s competence forlithic manufacturing, may provide rich information
with regard to the skill expressed materially by
the past knappers along with the operationalsequences in the lithic production. Indeed, some of
the Magdalenian open-air sites in the Paris Basin,
France, have offered well-known case studies tomake an attempt to address it (Bodu et al., 1990
and many others).
In Japan, there are a number of the refitted
materials recognized in the Upper Paleolithic sites.
Especially, the lithic refitted materials obtained
from the Shirataki sites, Eastern Hokkaido, areunique in the Upper Paleolithic record in Japan.
First, numerous refitted materials are recognizedfrom each of the assemblages, having a wealthof technological information concerning the
reductions of blades, microblades, bifaces, and boat-

shaped tools respectively. It apparently enables us
to compare the refitted materials within particularassemblages supposed to be an identical cultural
tradition. The procedure of such comparison may
prove a useful way of considering the unevenly
expressed skill within certain assemblage, inorder to explain the learning process among thepast knappers. Second, several obsidian refitted
materials, conjoining with many of stone tools anddebitage, sometimes reconstructed nearly original
raw material, are recognized. These materials cancontribute to surmise the morphological featuresof gravel and lithic debris used as original raw
material in the respective assemblages. Last,
we can assess the spatial distributions of therefitted lithic artifacts within the respective sites,
although these were to some extent modified
by natural transformation processes such as
solifluction. Analyzing these distributions is
critical to answering the questions concerning
the skill learning processes, highlighting how the
distributions of the analyzed refitted materials
relate to the prehistoric human behaviors occurred
within site.
In this paper, I attempt to analyze the refitted
materials obtained from the Kyushirataki 15 site,and discuss two topics mentioned below.
1) How the operational sequences of the pastknapping were segmented in the progress ofmanufacturing and organized? Also, how can weunderstand it?
2) Did a shift of the knapper in the progress ofoperations happen? Also, how can we identify it?

Figure 1. Refitted material No.96 from the Kyushirataki
15 site. This shows that large debris of obsidian was usedas raw material for blade reduction. The former stage ofmanufacturing was aiming to detach the large blades,with careful preparations and rejuvenations of core. In
the progress of blade manufacturing, large blocks were
gained by splitting and used as blanks for manufacturingof the boat shaped tools. After that, the small blades weredetached from the diminished core again. These resulted
in face battering and stacked steps on the flaked surface
and platform of the core.

Learning Behaviors in Prehistoric and Modern Humans

Figure 2. Artifacts included in the refitted material
No.96. The Manufacturing stage of this refitted material
can be divided from 1 to 34 (Naoe, 2012). 1-2: blades
produced from the manufacturing stage 12, 3: blade
produced from the manufacturing stage 14, 4: the boat
shaped tool produced from the manufacturing stage 15,
5: blade produced from the manufacturing stage 23, 6:
blade produced from the manufacturing stage 25, 7:
discarded blade core049

“Gift” as a Means of Cultural Transmission: An
Archaeological Implication of Bow and Arrow
Technology in Papua New Guinea
Yoshihiro Nishiaki

The University Museum, The University of Tokyo, Japan

Learning Behaviors in Prehistoric and Modern Humans

050

Introduction
A primary goal of the research project RNMH
is to investigate how the learning behaviors of
Neanderthals and modern humans differed inregard to their archaeological records. The evidence
currently being evaluated includes a combinationof the material remains recovered fromarchaeological sites and the data obtained throughapproaches known as experimental archaeology
and ethnoarchaeology.
I n t h i s p a p e r, I w i l l d i s c u s s a n
ethnoarchaeological approach that explores how
observations related to a modern human society
can be utilized to understand prehistoric learning
behaviors. A case study will be presented using datacollected in 1971 by Hitoshi Watanabe (1919–
1998) on the bows and arrows belonging to ahunter-farmer’s community in Papua New Guinea.
Watanabe’s innovative fieldwork strategy, named byhim as the “bow-and-arrow census,” documented
detailed morphological data on all of the existingbows and arrows in the study settlement alongwith personal information about the individuals
who manufactured and owned them. This uniquecorpus of data provides us with a rare opportunity
to investigate the learning processes involved in
bow and arrow manufacturing and the use of thistechnology in a pre-industrial society.
Watanabe’s bow and arrow census data from
Papua New Guinea
Watanabe’s fieldwork was carried out at thenative village of Wonie in western Papua New

Guinea between July and August 1971. The village
consisted of 16 houses/families, with a populationof 103, including infants. The primary formsof subsistence of the village’s inhabitants were
hunting and farming (with men doing the hunting
and women the farming); hence, they were hunterfarmers, or hunter-horticulturalists.
The uniqueness of Watanabe’s research lies in
its methodology. He collected census data on the
population and the bows and arrows present in
the village. The census data offer comprehensive
information on sex, age, education, occupation,family, experience of hunting, etc., whereas thedata on the bows and arrows include raw material,measurements, manufacturer, owner, and so on.
Most of these data are presented in Watanabe’smonograph (1975) and additional data can beobtained from the Hitoshi Watanabe collectionthat was donated to the University Museum,
Tokyo, after his death (Nishiaki 2007, 2011). The
latter collection includes original specimens suchas arrows and bows, manufacturing tools, toy bows
and arrows, video tapes, photos, and numerous
written records.
Analysis
My research goal is to establish a model on thelearning process behind bow and arrow technology.
This model incorporates information on how anindividual learns to make and use the bow andarrow, during his lifetime and how that knowledge
is transmitted from one generation to the next.
Data on the bows and arrows were analyzed inrelation to the age of their manufacturers andowners. Particular attention was paid to theownership of each bow and arrow, since Watanabeemphasized the tradition of giving bows andarrows in the Wonie society. In addition, changes
in the technological patterns of the bow and arrowby age group (e.g., infant, child, adolescent, adult,
and elderly) were considered:
Using: what type of bow and arrow is owned byeach age group; and what type of game is hunted
by each age group.
Manufacturing: which age group makes whichkind of bow and arrow; and the level of the
manufacturing skill expressed by each age group.
Giving : the age group of both giver and recipient
and the type of bow and arrow given.

Implications for prehistoric archaeology
Watanabe noted the importance of giving orgift in cultural, sociological, and ecological terms.
I would like to emphasize its importance in theteaching-learning system of a society. As a matterof fact, giving seems to play a primary role in thetransmission of bow and arrow technology inWonie society. While the motive behind giving
was not specified in Watanabe’s records, there wasno obvious reciprocation between the giver andrecipient. The scenario is probably comparable towhat Mauss (1990) described on primitive society’s
giving tradition in the early 20th century. The giftof bows and arrows was made by the elders to theyoung individuals, who then reciprocated when
they became older. This system cycled throughout
a generation.
This gift-education model can serve as a guidefor exploring differences in the learning behaviors
between Neanderthals and modern humans
and their consequences on the archaeologicalrecord. For example, the domain-specificprogress in learning can be tested with carefulanalyses of the techno-typological features of
the archaeological objects. In addition, the life
histories of Neanderthals and modern humans canbe examined from a biological point of view. Was
adolescence in Neanderthals shorter in duration
than in modern humans? Finally, was there any
evidence of giving among Neanderthal societies?
We have a lot of evidence of trade and perhaps
also giving for the modern human sites, but littleincontestable evidence exists in the archaeologicalrecord of the Neanderthals. If adolescence was

Learning Behaviors in Prehistoric and Modern Humans

Results
The analysis yielded the following observations.
First, the learning of bow and arrow technology isdomain-specific and it progresses cumulatively. The
knowledge and skill of the manufacture and use ofthis technology, as well as the decorative style, do
not develop simultaneously. For example, infants
only use the bow and arrow during play, whilechildren use them for capturing small animals
such as birds and fish. Later in their childhood,they begin manufacturing their own bows. The
manufacture improves during their adolescence,
and when they reach early adulthood, all of thenecessary elements involved in bow manufacture,use, and decoration are established.
Second, learning bow and arrow technology
is part of a dynamic system. As in most huntergatherer societies, there is no evidence of teaching.
However, the Wonie case suggests that theirtradition of giving serves as a means of teaching.
In nearly all cases, bows and arrows are given
by older individuals to younger ones, with a few
exceptions among the adult group, who exchanged
products with one another. Younger individuals are
given bows and arrows specific to their age group.
When younger individuals age, they manufacture
the same type of bow and arrow that they received

early in life and then give these items to theyounger age groups. The knowledge of bows andarrows thus continues through cycles of giving.
Third, knowledge and technological skills of thebow and arrow differ among age groups. The mostdistinctive period involves adolescence, when thetypes of bows and arrows given to individuals vary
the most. This suggests that adolescent individuals
learn more during this period of their lives thanthe other age groups.

051

indeed shorter and there was no establishedtradition of giving, the learning of material culture
in Neanderthal society likely differed significantly
from that of modern humans as well as modern
hunter-gatherers, such as the Wonie of Papua New
Guinea.
References
Mauss, M. (1990) (1922) The Gift: Forms and functions of
exchange in archaic societies. London: Routledge.
Nishiaki, Y. (2007) Catalogue of Anthropological Materials
in the Hitoshi Watanabe Collection. The University
Museum, The University of Tokyo, Material Reports
68. Tokyo: The University of Tokyo.
Nishiaki, Y. (2011) An ethno-archaeological approach tothe study on prehistoric learning. In: Proceedings ofthe 4th Conference on the Replacement of Neanderthals
by Modern Humans, edited by C. Tanabe, pp. 44–45.
Okazaki: Okazaki Conference Center.
Watanabe, H. (1975) Bow and Arrow Census in a West
Papuan Lowland Community: A Field for Functionalecologic al St udy. Universit y of Q ueensland,
Anthropological Museum, Occasional Papers
in Anthropolog y 5. Auckland: University of
Queensland.

Learning Behaviors in Prehistoric and Modern Humans

052

The Effects of Cross-Boundary Rituals on Cultural
Innovation

Shiro Horiuchi1 and Sachiko Kubota2

1Shibaura Institute of Technology, Japan 2Graduate School of Intercultural Studies, Kobe University, Japan

Introduction
In this talk, we attempt to test the hypothesis thatthe cross-boundary rituals, in which members ofdifferent groups communicate with one another,affected the development of the culture of modern
humans. To test this hypothesis, we introduce anagent-based model (ABM) simulation. We thenintroduce case studies of residents of rural areas inJapan and Australian Aboriginal people to show
how present-day modern humans performed crossboundary rituals.

Figure 1. If agents migrate between groups (x > 0) andagents of many cultural traits lead cross-boundary ritual (0< y < ∞) ,the average value of Σ c increases significantly.

agents copy others’ cultural traits within their groups
as well as at cross-boundary rituals.
The simulations suggest that ∑ j Ci,j swiftly increases
if the cross-boundary rituals are held between agentsof relatively many cultural traits and any agents
migrate between different groups (Figure 1). The
modern humans should have performed such crossboundary rituals and migrated between groups toexpand their advanced culture.

Japanese rural ritualTraditional rituals, “spirit dances” or “kagura”,
continue to be performed in many rural areas ofJapan. Recently, an increasing number of tourists
visit kagura performances as audience members andthe kagura performances have become the crossboundary rituals. Some kagura performances arestill within-boundary rituals in that kagura limit
participation to local residents. Thus, to test the effectsof cross-boundary rituals on present-day modern
humans, we can compare kagura groups in terms of
how they deal with tourists. The questionnaires were
mailed to 155 leaders of kagura groups in Miyazaki
prefecture; 87 (55%) of whom responded. Audiencesfor 49 groups were composed primarily of local
residents (within-boundary rituals) and those for 39
groups consisted primarily of tourists (cross-boundary
rituals). The number of dancers and audiences
increased in 55 (active) groups and decreased in 32(non-active) groups, respectively, for these 10 years.
The data suggest that in within-boundary rituals,
members should limit participants to local residents

Learning Behaviors in Prehistoric and Modern Humans

ABM simulation
To test what types of rituals are necessary for culturalaccumulation, we build an Agent-Based-Model(ABM) simulation. The ABM assumes N agents andM groups. Culture consists of K independent traits,
represented as the vector (Ci,1, Ci,2, …, Ci,K) for agent i.
If the agent knows or does not know the j th cultural

trait, Ci,j is 1 or 0, respectively. In the simulation,053

and recruit from their communities to activate theirkagura. In cross-boundary ritual, in contrast, members
should communicate with tourists and advertise
their local communities. Furthermore, leaders ofkagura performing cross-boundary rituals found thatthey could educate young dancers and experiencekagura as a form of enjoyment in activated kagura.
Accordingly, cross-boundary rituals affect the culturalaccumulation of contemporary modern humans
by enlivening kagura through communication
with outsiders. Such mechanisms were likely to be
operating during the age in which the replacement
occurred.

Learning Behaviors in Prehistoric and Modern Humans

054

Australian Aboriginal ritual
In northeastern Arnhem Land, in the Northern
Territory the Yolngu language speakers live infour villages. The Yolngu still value many of theirtraditional practice especially their rituals. Theserituals remain at the core of their social life andpeople devote considerable energy to them. Besides
localized rituals, there are larger ritual festivals that
not only bring members of many Yolngu clans
together but people from other linguistic groups. On
these occasions people may exchange their dancesand songs. Through such festivals groups of distantly
related people make alliances and trade.
There was the opportunity for wife exchange insome ceremonies where visitors came from far away.
The organizers of a Kunapipi ceremony would send a
messenger informing other groups that they planned
to hold a ceremony. To do it in the correct way, theritual intercourse had to be carried out with theconsent of the female partner, which was essential
if it was of not to cause conflict. Men intending toswap wives had to send presents via a messenger
who was in a special relationship to the receiver. On
the last day of the ritual, after the previous exchangeof messages and gifts, the men got together at thecamp site of the Kunapipi ceremony and sat in acircle, with women dancing in the distance, showing
their willingness to accept intercourse by a feathered
headband on their forehead.
Currently in Arnhem Land, those old sexual
practices have been abandoned, although, Kunapipi

rituals remain a focus of social life. So in thecontemporary situation, the ritual still functions as anoccasion for the transmission of traditional knowledge
from old to young, and as an important occasion
for socializing and exchanging of knowledge withpeople from distant places. And today because more
people can come together by plane, boats and cars, it
increases the opportunity of active cultural exchange.
Conclusion
Durkheim depicts the function of rituals as theintensification of sense of solidarity of the group.
But as we have seen, ritual festivals are not attended
only by the local population but are also attended
by people from far away who do not interact ona frequent basis. The exchanges during the rituals
opened up opportunities for knowledge exchange
and were probably important sources of cultural
innovation. They could also lead to intermarriage,
migration and genetic exchange which increased thechances for stable cultural innovations.

Teaching, Trust, and Early Rapid Social Learning in
Hunter-Gatherers

Barry S. Hewlett

Washington State University, USA

The cognitive scientists also indicate that theability to read the intentions of others (theory ofmind) is not necessary for natural pedagogy to
evolve, but that learning from natural pedagogy ismuch more likely to be efficient, early, and rapid if
the learner trusts and feels secure with the teacher.
The paper describes the contexts of infant and childdevelopment among Aka hunter-gatherers that
promote trust and empathy in child development.
The paper emphasizes how interactions between
inherent evolved cognitive mechanisms, suchas natural pedagogy, and contemporary huntergatherers’ culturally constructed environments,
amplified social learning and made cumulative
culture a distinctive feature of modern Homosapiens.

Learning Behaviors in Prehistoric and Modern Humans

Prominent cultural anthropologists Margaret
Mead, David Lancy, and Barbara Rogoff indicatethat teaching does not exist or is rare in smallscale cultures. By contrast, recent research bycognitive neuroscientists Gyorgy Gergely andGergely Csibra indicate that one type of teaching,called natural pedagogy, is a human universal,
part of human nature, and not found in the great
apes. They hypothesize that this form of teaching
emerges in infancy and that it enhances humans’ability to faithfully transmit “opaque” cultural
knowledge, such the function of a particular tool.
Learners evolved to pay attention to particular
cues, such as eye and body movements, andteachers evolved the skills to convey important
information to learners, such as pointing, using
personal names, looking at or making sounds about
important knowledge. Cognitive science researchon natural pedagogy is limited because all of
their studies have been conducted in laboratories
with Western infants. This study uses videotapesof 10 Aka hunter-gatherer 12-14 month-old
infants in naturalistic settings to evaluate thenatural pedagogy hypothesis. The study showsthat natural pedagogy exists in hunter-gatherers,
but that it occurs relatively infrequently. The study
also identifies two new forms of hunter-gatherer
teaching that are rare or do not exist in the great
apes—distributed teaching and opportunity
scaffolding. These new forms of teaching occur
more frequently than natural pedagogy and may
have other social learning functions other thantransmitting opaque knowledge.

055Determinants of Cultural Evolutionary Rates

Kenichi Aoki

Graduate School of Science, The University of Tokyo, Japan

Human-Specific Learning Strategies and Cultural Evolution

056

The “learning hypothesis” seeks to accountfor the replacement of Neanderthals by modern
humans (Homo sapiens) in terms of an innate
difference in learning strategies (abilities) between
the two species. It comprises three proposals thatform a “syllogism”: 1. there is an innate differencein the learning strategies employed by modern
humans and Neanderthals; 2. the difference inlearning strategies translates into the observed
difference in cultural evolutionary rates and culturalcontent between the two species; 3. the differencein cultural evolutionary rates and cultural content
contributed significantly to the extinction of the
Neanderthals and their replacement by modern
humans.
Empirically, we know or suspect two things
that are relevant to proposal 3. First, the stone
tools made by modern humans, after say 50,000
years ago, underwent rapid changes compared tothose made by Neanderthals. That is, there was asignificant difference in the cultural evolutionary
rates exhibited by the two species during the
Paleolithic. Moreover, modern humans also had
ornaments, art, and tools made of materials otherthan stone. Second, replacements of modern
human ethnic groups by other modern humanethnic groups in historical times—notably theEuropean expansions into the Americas—were
driven by differences in the cultural content of thecompeting groups and by demographic differences
contingent on the differences in cultural content.
These observations suggest that, if Neanderthals
and modern humans did indeed compete, the moreculturally advanced modern humans would haveprevailed.
Our research group, B01, has as its goal thetheoretical justification of the learning hypothesis.
Evolutionary theory cannot be used to prove
proposal 1, i.e. the existence of innate differencesin learning strategies. Research groups C01 and
C02 will eventually provide archaeo-neurologicalevidence for size differences between modern
humans and Neanderthals in the areas of the brain
involved in individual learning and social learning.
Meanwhile, evolutionary theory can generatepredictions as to what kinds of learning strategies
will evolve under given environmental conditions,more specifically under given patterns of temporal
and/or spatial environmental variability. Then, in
combination with the empirical data provided
by research group B02 on the environmental
conditions experienced by modern humans andNeanderthals, which presumably were different, we
can plausibly argue for an innate difference in thelearning strategies of the two species.
The specific theoretical question that I ask in thispaper is: how does a learning strategy determine
the cultural evolutionary rate? That is, I will be
addressing proposal 2 of the learning hypothesis in
my talk, interpreting this as a statement in logic.
Before doing so, let me define what is meant by
a learning strategy. Its components are individual
learning (i.e. learning from personal experience, e.g.
by trial-and-error) and social learning (i.e. learningby copying others, e.g. by imitation). A learning
strategy is the way in which individual learning andevolution. For example, when long-term changes
in many cultural traits are under consideration, we
can adapt the infinite sites model of population
genetics, yielding the formula R =m∑ Niui π1ii=1

for the cultural evolutionary rate. Here, m is thenumber of social roles (e.g. male and female, ageclasses, expert and novice), Ni is the number ofindividuals in social role i , ui is the innovation
rate of an individual in social role i , andπ1i is the
fixation probability of an innovation made by anindividual in social role i .
One application of this formula is to the questionof whether one-to-many social transmission will
accelerate cultural evolution, as has been proposedby some archaeologists and anthropologists. Basedon a simple model, I show that one-to-many socialtransmission per se has no effect on the cultural
evolutionary rate.

Human-Specific Learning Strategies and Cultural Evolutionsocial learning are combined, either simultaneously
or sequentially, and the relative dependence on
each. The modes, pathways, and biases of social
learning are also integral components of a learning
strategy. Using these terms, a short definition ofcultural evolution is: the spread through social
learning of innovations produced by individual
learning.
In this paper I discuss two theoretical studies
showing that the innovation rate contributes moststrongly to the cultural evolutionary rate. Sincea higher rate of innovation requires an advanced
aptitude for individual learning, this argument
entails that Neanderthals were relatively deficient inindividual learning compared to modern humans,
whereas they were equally adept at social learning.
Other researchers have argued for the importanceof modes, pathways, and biases of social learning,such as “one-to-many” social transmission where
one expert serves as the teacher to many novices,and “direct bias” where the individual with the“best” variant of a cultural trait is identified andcopied. Demographic parameters such as the total
population size and the number of acquaintances
have also been invoked. The relevance of suchfactors is also an empirical question that needs
to be resolved by research groups A01 and A02.
Theoretically speaking, however, they are not asimportant as the innovation rate in determining
the cultural evolutionary rate.
Definition and anal y sis of the cultural
evolutionary rate utilizes the analogies between
genetic (i.e. biological) evolution and cultural057

Scheduling of Individual and Social Learning as
Optimal Life History Strategy

Joe Yuichiro Wakano1 in collaboration with
Kenichi Aoki2 and Laurent Lehmann3

1Meiji University, Japan 2The University of Tokyo, Japan 3University of Lausanne, Switzerland

Human-Specific Learning Strategies and Cultural Evolution

058

In the replacement of Neanderthals by modern
humans, innovative stone tools produced bymodern humans might have played an importantrole. Advanced and complex industries cannever be invented by a single individual without
learning from other individuals. They are results ofcumulative cultural evolution in which knowledge
is inherited from parental generations to offspring
generations. However, if all individuals simply
copied what had been already known, there would
be no advancement in technology. Contribution to
the culture by improving the preceding knowledge,
such as a discovery of new adaptive use or formof stone tools, is also crucial for cultural evolution.
Inheritance of culture is realized by social learningand improvement is realized by individual learning.
Thus, it is an important problem when the
cumulative culture develops fastest under different
learning strategies: when and how should anindividual perform learning in one's life?
The balance between a weight assigned tolearning and a weight assigned to the exploitationof the learned knowledge is also an interesting
problem. To maximize the accumulation rateof culture, one should spend sufficient time toabsorb the existing knowledge in the parent
generation and then spends the rest of one's
lifetime to improve the knowledge by individual
learning. However, such a learning schedule does
not necessarily maximize the fitness of the focal
individual, i.e., the expected number of biologicaloffspring. Possible scheduling of social learning,
individual learning, and the exploitation of theknowledge is illustrated in Figure 1.
These types of problems have been studied in
the field of behavioral ecology and referred to as
the optimization of life history strategy. Several
mathematical frameworks such as Pontryagin's
maximum principle or dynamic programmingare known. A famous example is the optimalschedule of allocation of resource into growth andreproduction in plant's life history.
Recently, some researchers have applied theseframeworks (optimal scheduling) to study cultural
evolution. Conceptually speaking, this question
had been already considered in the model of Social
Learning Tournament. After a brief (and nonmathematical) overview of the previous studies,
we will introduce some of our recent results on
optimal learning schedule based on mathematicalmodeling and analysis. We find that social learningand individual learning occurs in this order inthe evolutionarily stable schedule. This order isconsistent with the one necessary for cumulative
cultural evolution (because otherwise modification
or improvement to pre-existing knowledge does
not occur.) We also find that the maximizationof personal fitness greatly contradicts the
maximization of cultural level of a whole
population. That is, even though much higher levelof culture can be sustained if people spend somemore time in learning, the competition between
individuals does not allow a population to reachthis level. The situation is somehow similar to a
public goods game where public goods represent
the level of culture of a whole population. Our

results provide a basis of a working hypothesisthat the difference in lithic cultures between
Neanderthals and modern humans comes from thedifference in learning schedules.
Figure 1. An illustration of a learning schedule. Three
discrete generations are shown. The duration of infant/juvenile/adult periods determines the accumulation rateof culture, the equilibrium level of culture, and personal
biological fitness.

Human-Specific Learning Strategies and Cultural Evolution

059

The Handaxe and the Microscope: Experience-Basedand Fictitious Learning in a Geometric Model of
Adaptation
Laurent Lehmann

University of Lausanne, Switzerland

When individuals learn by trial-and-error, they
perform and experience randomly chosen actions
and reinforce those actions providing high payoffs.
Cognitively more sophisticated individuals may
evaluate the consequences of actions throughmental simulation without experiencing themphysically.
Such fictitious learners can select actions withhigh payoffs without making long chains of trialand-error learning. Here, we analyze the evolutionof an n-dimensional cultural trait (or artifact)
under experience-based and fictitious learning in apayoff landscape with a single optimum. We derive
the stochastic learning dynamics of the distance tothe optimum in trait space when choice between
alternative artifacts follows the standard logit
Human-Specific Learning Strategies and Cultural Evolution

060

choice rule. We show that for experience-based andfictitious learners, the learning dynamics stabilizes
away at approximate distance [n/(2L)]^(1/2) fromthe optimum, where L is an effective learning
performance parameter depending on the learning
rule under scrutiny. Learners, whether simple
or sophisticated, are thus unlikely to reach the
optimum when traits are complex (n large) andthus face a barrier to improvement. We showthat this barrier can be significantly reduced in alarge population of learners performing payoffbiased social learning, in which case L becomes
proportional to population size. Overall, our results
illustrate the effects of errors in learning, levels ofcognition, and population size in the evolution ofcomplex cultural traits.

Social Learning, Trial-and-Error, and Creativity

Nobuyuki Takahashi, Ayaka Hatano, Misato Inaba, Ryoichi Onoda,
and Dora Simunovic
Center for Experimental Research in Social Sciences, Graduate School of Letters, Hokkaido University, Japan

These previous studies have two problems,
however. First, there is little empirical data on therelationship between individual learning and social
learning among humans. Second, they assume thatsocial learning is usually represented by the single
ability of imitation, while individual learning isrepresented by either trial-and-error or creativity,
although these two abilities are quite different
from each other. Therefore, in the current study, weinvestigated the relationships between these threemajor human learning abilities for the first time.
We have conducted two experiments. Bothexperiments were conducted in the Group
Experiment Lab at CERSS, Hokkaido University.
Participants were asked to perform tasks designed
to measure trial-and-error, creativity, imitation,and certain other abilities. They also answereda questionnaire which asked various questions
regarding learning strategies.
Fifty-eight undergraduate students participatedin the first experiment. The first experimentconsisted of three types of tasks. The first one
measured trial-and-error.We used the Lonpos gamein which participants were asked to fill in empty
spaces on a board with different-shaped pieces.
The second task measured creativity. For that weused the two-string problem (Maier, 1945) andthe candle holder problem (Dunker, 1945) both ofwhich are known to measure functional fixedness
in cognitive psychology. The third task measured
imitation. For that we used a wire puzzle to see if
participants become capable of solving the puzzle
after watching a video of a demonstrator solving the

Human-Specific Learning Strategies and Cultural Evolution

Homo sapiens is a species whose behaviors
are determined heavily by learning. It has been
speculated that Homo sapiens have acquired sucha high level of learning ability because it has
been adaptive during the evolutionary history.
Research on learning that employ mathematicalmodeling have assumed implicitly a dichotomy
between individual learning and social learning,and examined under what conditions each type oflearning ability can be adaptive. The key difference
between these two is whether the source of
information originates in oneself or others. Usuallyindividual learning is considered a more basic
ability since species who have very limited cognitive
abilities, such as Paramecium caudatum, can have it,whereas social learning is a more advanced ability
since it requires sophisticated cognitive and mental
abilities.
One of the key insights in previous research is thatcost and benefit of individual and social learning
abilities are different. For an individual to learn byhimself, he has to invest his resources. However,other individuals can imitate the fruit of his labor at
no extra cost to themselves. This presents us with afree-rider problem. For each individual, it is betterto engage in social learning. However, if everyone
engages in social learning, new knowledge would
not be acquired. Therefore, many previous studies
assumed that there is a tradeoff between individual
learning and social learning, and examined underwhat conditions social learning can be adaptive (e.g.,
Borenstein et al. 2008; Kameda & Nakanishi 2002;
Wakano et al. 2004)061

Human-Specific Learning Strategies and Cultural Evolution

062

puzzle. The results suggested that the relationshipbetween creativity and trial-and-error was negative,
although they are usually both considered aspects ofindividual learning. The results also suggested that
there was no trade-off between individual learningand social learning since no ability was negatively
related to imitation. Finally, only creativity was
positively related to imitation, which partly supports
a new theoretical argument that a combinationof social learning and individual learning is mostadaptive under certain fluctuating environments(e.g., Aoki, 2010).
However, what we found in the first experimentis far from conclusive evidence since it was
obtained from only a few specific tasks. Therefore,
we conducted the second experiment to see if
we can conceptually replicate the result of thefirst experiment. Sixty undergraduate students
participated in the second experiment. In it, we useda new task to measure trial-and-error. We developeda coin game in which three buttons: a circle, triangle
and a star, were displayed on the participant’s
computer screens. If participants clicked the
buttons in a certain way, they would be rewarded
with coins on the computer. Their objective wasto figure out what pattern of clicking the buttons
would get the coins. This is conceptually equivalent
to the Skinner box, except that the reward is notreal food. For creativity, we used several new tasks
in addition to the candle holder problem. First, weused Cattell CFIT scale 3 form A, which is a testdeveloped to measure fluid intelligence. Second,
we used the ring task (McCaffrey, 2012). Third, weused two paper-and-pencil tasks; the truck task andthe light task (McCaffrey, 2012). For imitation, weused the same wire puzzle. The results suggested
that, contrary to the result of the first experiment,
the relationship between creativity and trialand-error was positive. There was no correlation
between trial-and-error and imitation, same as in
the first experiment. Finally, creativity was positively
related to imitation, as was also the case in the firstexperiment. In conclusion, the results of the second
experiment replicated most of the ones from thefirst experiment, except for the relationship between

creativity and trial-and-error. The candle holder
problem was used in both experiments, yet it was
negatively related to the Lonpos task and positively
related to the coin game. The next step is to examine
the validity of these two tasks.

Experimental Studies of Modern Human Socialand Individual Learning in an ArchaeologicalContext: People Behave Adaptively, but within Limits
Alex Mesoudi

Department of Anthropology and Centre for the Coevolution of Biology and Culture
Durham University, UK

technology in general [3-5]. Participants in small
groups of 5-6 each design an arrowhead via a
computer program (Figure 1). This virtual arrowhead
is composed of three continuous traits (Height,
Width, Thickness), which can each take any value
from 1-100 arbitrary units, and two discrete traits(Shape and Colour), which can each take one offour categorical values. Over a series of trials (or‘hunts’), participants can improve their arrowhead byeither individual trial-and-error learning, by directly
altering the traits, or social learning, by copying thedesign of another group member. The form of thissocial learning (e.g. payoff bias, conformity) can be
manipulated. On each hunt the participant tests
their arrowhead in a virtual hunting environment,
receiving a score in calories: the closer their design isto one or more hidden optimal designs pre-specified
by us using fitness functions, the higher the score.
Of particular interest is when there are multiple
locally optimal designs, giving a realistic multimodal

Figure 1. A screenshot of the Virtual Arrowhead
Task. Participants can choose to directly change
the traits in the box at the top (individual learning)or copy the design of another participant in the box
on the left. Feedback is given in calories depending
on how close the design is to one or more hidden
optimal designs

Human-Specific Learning Strategies and Cultural Evolution

The aim of this conference is to explore thehypothesis that a key factor in the replacement of
Neanderthals by modern humans was the superiorlearning abilities of the latter. To properly evaluate
this hypothesis, empirical data are needed on thelearning abilities of modern humans. This paper
reviews a set of experimental studies conductedby myself and collaborators that have probed thelearning abilities of contemporary humans when
faced with a novel and complex task designed toresemble technology found in the material record.
Hopefully, the findings of these experiments caninform both theoretical models of Neanderthal
replacement and interpretation of the often
ambiguous archaeological record.
The Virtual Arrowhead Task was originallydesigned by myself and archaeologist Michael
O’Brien to capture the key aspects of North
American projectile points [1,2], although we havesince used it to explore the learning of complex

063

adaptive landscape, implemented by giving thecontinuous traits bimodal fitness functions. Color istypically neutral and does not contribute to fitness
in any way, and there is always a small random error
in the score, simulating stochastic conditions such as
weather or prey availability.
This task is intended to capture the key aspects ofmost complex technology, including that used byboth modern humans and Neanderthals around thetime of their coexistence: a technology composedof multiple constituent traits (some continuous andsome discrete, some functional and some neutral),that is functionally opaque (no obvious, intuitive
relation between an artifact and its effectiveness) andwhich has multiple locally optimal alternative designs(i.e. a multimodal adaptive landscape). In a series ofstudies we have explored how contemporary humans
engage with this task, with the following key findings:

Human-Specific Learning Strategies and Cultural Evolution064

1. People are effective individual learners, but can
get stuck on local optima
When playing alone, participants engage in a simple
but effective reinforcement learning strategy [1,2]:
modify a trait (e.g. increase Width), if the payoff
increases then keep modifying the trait in that way(e.g. increase Width further); if the payoff decreases
then do the opposite (e.g. decrease Width). This isdone for each trait in turn until payoffs no longer
change. The magnitude of the change negatively
correlates with the participant’s current payoff, i.e.
low payoffs provoke bigger changes. This is adaptive:it allows big jumps across the adaptive landscapefrom low-fitness valleys to potentially high-fitness
peaks. The downside of individual learning is that,in multimodal adaptive landscapes, participants getstuck on locally optimal but globally sub-optimal
peaks.
2. People use payoff-biased social learning to jump
to higher-fitness designs
When social learning is allowed, either after aperiod of individual learning [1] or concurrently
with individual learning [4], participants readily
engage in payoff-biased social learning, copying thedesign of the most successful person in their group.
This is adaptive, as it allows them to outperform

pure individual learners by jumping from lowfitness local optima to a higher-fitness peak thathas been found by a better-performing participant.
This use of social learning is flexible and dependent
on the participants’ own performance: only when
performing poorly do participants engage in payoff
biased social learning.
3. Social learning is used more when individual
learning is costly
When individual learning is made more costly,
by deducting calories every time a trait is directly
modified, participants respond adaptively byengaging in social learning more frequently [1].
4. Payoff biased social learning is preferred toother forms of social learning
Payoff bias is preferred to conformity (copying
the most common design), averaging (copying
the average values of each trait in the group) andrandom copying (copying a randomly-selectedgroup member) [5]. This is adaptive, as only payoffbias allows participants to locate high-fitness peaks.
5. Payoff biased social learning leads to “cultural
hitchhiking”
As noted above, Color was neutral and had no effecton payoff. Yet Color was copied just as faithfully
as the functional traits during payoff-biased social
learning [1]. This is an example of a neutral trait
hitchhiking on functional traits; while this trait was
neutral, it is possible that maladaptive traits also
hitchhike in this way.
6. Informational access costs block social learning
In the above, participants could freely viewother participants’ designs. In one study, we letparticipants set ‘informational access costs’, which
potential copiers had to pay them in order to view
their arrowhead designs [4]. This blocked virtually
all social learning, as the most successful participants
set unreasonably high access costs that no otherparticipant was willing to pay.
7. Separating individual learners from participants
inhibits social learning
In one study, a separate pool of individual learners
was used, rather than allowing participants to copy
fellow group members [5]. As before, payoff-biased
social learning was objectively the best strategy touse, yet only a minority of participants engaged in it,
possibly due to the reduced salience of the models.
8. Looking times are just as effective copying cues
as objective performance
Participants preferentially copy other participants
whom they have been told have been viewed longer
by other group members [3]. This looking time
information, a potential marker of prestige, was usedjust as often as objective success, suggesting that
prestige may be a common shortcut for identifying
who to copy.
Conclusions:Contemporary humans behave largely adaptively
when faced with a relatively complex and novel
technology-design task, using feedback on theirperformance to flexibly switch between individual
learning and payoff-biased social learning, andeschewing non-adaptive social learning strategies
such as conformity and random copying. However,there are limits to this adaptiveness: information is
not shared freely, at least in competitive tasks like
this; neutral traits are unnecessarily copied alongside
functional traits; separate pools of individual learners
are not exploited well; and looking time information
is used to identify from whom to copy as often as
more reliable measures of objective success.
Human-Specific Learning Strategies and Cultural Evolution

References: [1] Mesoudi & O'Brien (2008) The culturaltransmission of Great Basin projectile point technology
I: An experimental simulation. Am. Antiq. 73, 3-28. [2]Mesoudi & O'Brien (2008) The cultural transmissionof Great Basin projectile point technology II: An
agent-based computer simulation. Am. Antiq. 73, 627644. [3] Atkisson, Mesoudi & O’Brien (2012) Adult
learners in a novel environment use prestige-biased social
learning. Evol. Psychol. 10, 519-537. [4] Mesoudi (2008)
An experimental simulation of the 'copy-successfulindividuals' cultural learning strategy. Evol. Hum. Behav.
29, 350-363. [5] Mesoudi (2011) An experimental
comparison of human social learning strategies. Evol.
Hum. Behav. 32, 334-342.

065

Digital Reconstruction of the Neanderthal Amud 1
Cranium
Naomichi Ogihara1, Takeo Kikuchi1, Yusuke Morita1, Hideki Amano1,
Osamu Kondo2, Masato Nakatsukasa3, Hiromasa Suzuki4, Takashi Michikawa4,
Hajime Ishida5, and Takeru Akazawa6

1Department of Mechanical Engineering, Faculty of Science and Technology, Keio University, Japan 2Graduate School of Science, The Universityof Tokyo, Japan 3Department of Zoology, Graduate School of Science, Kyoto University, Japan 4Research Center for Advanced Science andTechnology, The University of Tokyo, Japan 5Faculty of Medicine, University of the Ryukyus, Japan 6Kochi Institute of Technology, Japan

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

066

Fossil crania are often fractured and fragmented
due to compaction and diagenesis. To restore theantemortem appearance of a fossil cranium, itis therefore necessary to correctly assemble thefragments. Conventionally, such reconstructions
are created manually based on the knowledge andexperience of skilled anthropologists. However, toinvestigate the difference in learning ability between
Neanderthals and early modern humans in terms ofanatomical differences in their brain morphology, it
is essential to develop new computerized methods torealize more objective restorations of fossil crania. In
this study, we developed a computerized techniquefor assembling fossil cranial fragments based onshape information, i.e., the smoothness of the joints
between fragments and applied this technique forreconstruction of the Neanderthal cranium Amud 1.
We virtually removed glue and plaster fromthe original specimen and separated off thefragments constituting the fossil cranium. We
then approximated the surface of each of cranial
fragments using Bezier surface to extrapolate
the surface and mathematically predict shape ofadjacent fragments. The positions and orientationsof adjacent fragments were calculated one by one
by minimizing the fitting error. We assembled rightand left sides of the cranium separately based on theabove mentioned technique, and then put the rightand left parts together mathematically based onsmoothness of the joint, bilateral symmetry and the
distance between the mandibular fossae determinedfrom the associated undeformed mandible. As aresult, smooth yet globally consistent assembly of
the fragments of Amud 1 cranium became possible(Figure 1). The position of the isolated maxilla wasdetermined by its complete mandible articulated tothe mandibular fossae.
A reference modern human cranium was thenwarped onto the new reconstruction by iterativethin-plate spline deformation to restore complete
exso- and endocranial morphology. Here we utilized
the Gibraltar 1 to estimate position of landmarkson the missing basicranial and damaged endocranial
regions.
The comparisons of the present computerizedreconstruction with the original reconstruction
indicated that the newly reconstructed Amud 1
cranium is slightly shorter, wider and lower in theanteroposterior, mediolateral, and superoinferior
directions, respectively. Such computerized method
allows detailed documentations of how the cranium
was reconstructed, making restoration process
reproducible and verifiable, while eliminating someof the subjectivity that may occur in restoration.
To estimate brain morphology of Amud 1
based on the reconstructed fossil cranium, it isnecessary to quantitatively establish morphologicalcorrespondence between cranial endocast andbrain surface morphology based on living humansubjects using medical imaging technique. For this,the CT and MRI images from the same subjects
were acquired and registered to each other suchas to maximize mutual information between thetwo images. Endocast and brain surfaces were thenFigure 1. New assembly of the fragments of Amud 1
cranium.

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

three-dimensionally reconstructed, and landmark
coordinates were acquired to evaluate morphologicalcorrespondence between them. Our preliminary
analysis demonstrated that such morphologicalcorrespondence between cranial endocast and brain
surface does actually exist, suggesting that thiscorrespondence relationship might be exploited for
estimation of brain morphology of fossil humancrania. We plan to derive spatial deformation
functions from modern human crania to thereconstructed fossil cranium using thin-plate spline
function, and use them to deform human brains toestimate brain morphology of the fossil cranium.

067

Functional Craniology, Human Evolution, andAnatomical Constraints in the Neanderthal Braincase

Emiliano Bruner

Centro Nacional de Investigación sobre la Evolución Humana, Burgos, Spain

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

068

The relationships between face, cranial base, andneurocranium, have received much attention in thelast decade, mostly taking into account changes
associated with hominid evolution (Bookstein
et al., 2003; Bastir et al., 2006). In particular,
recent advances in digital anatomy and computed
morphometrics have allowed to go beyond theanalysis of individual traits variation, supporting
quantitative approaches to study the integration
between brain and braincase (Moss and Young,
1960; Richtsmeier et al., 2006). Neanderthals
display some derived endocranial features, namely
a relative widening of the prefrontal (Bruner
and Holloway, 2010) and upper parietal areas(Bruner et al., 2003). Nonetheless, the braincase
in Neanderthals retains more plesiomorphic
characters than the modern human neurocranium,
despite the similar cranial capacity in these two
taxa (Gunz et al., 2010). The general allometricpattern associated with brain size increase in thehuman genus involves a longitudinal constraint of
the midsagittal parietal surface, probably becauseof the contiguity with the frontal and occipital
volumes and because of allometric relationships
between cortical volumes and the falx cerebri(Bruner, 2004; Bruner et al., 2011a). In termsof bones, there is a marked integration between
parietal and occipital structure (Gunz andHarvati, 2007). Interestingly, Neanderthals display
hypostatic traits at the parieto-occipital junction,suggesting ossification problems associated withmorphogenesis and balance between growthand development (Sergi, 1934; Manzi, 2003;

Manzi et al., 1996). Neanderthals also lacked acomplex meningeal vascular network, opening to
speculations on possible thermal limits based onblood flow of the cortical surface (Bruner et al.,2011b). Taking into account all this information,
we must consider the hypothesis that Neanderthals
may have experienced structural problems in theirbraincase, reaching physical limits because of theplesiomophic neurocranial structure associatedwith a highly derived brain size. It is necessary toinvestigate whether or not such constraints may
have influenced the evolution of the Neanderthals,
their extinction, and their replacement by modern
humans. In this sense also the relationships
between structural organization and cognitive
functions must be carefully considered, evaluating
hypotheses to identify primar y causes andsecondary consequences.
References
Bastir M, Rosas A, O’Higgins P. 2006. Craniofacial levels
and the morphological maturation of the humanskull. J Anat 209: 637-654.
Bookstein FL, Gunz P, Mitteroecker P, Prossinger H,
Schaefer K, Seidler H. 2003. Cranial integration in
Homo: singular warps analysis of the midsagittal
plane in ontogeny and evolution. J Hum Evol 44:
167-87.
Bruner E. 2004. Geometric morphometrics andpaleoneurology: brain shape evolution in the genus
Homo. J Hum Evol 47: 279-303.
Bruner E, Holloway RL. 2010. A bivariate approach to
the widening of the frontal lobes in the genus Homo.
J Hum Evol 58: 138-146.

Manzi G. 2003. ‘‘Epigenetic’’ cranial traits, Neandertals
and the origin of Homo sapiens. Riv Antropol 81: 5768.
Manzi G, Vienna A, Hauser G. 1996. Developmental
stress and cranial hypostosis by epigenetic trait
occurrence and distribution: an exploratory study on
the Italian Neandertals. J Hum Evol 30: 511-527.
Moss ML, Young RW. 1960. A functional approach tocraniology. Am J Phys Anthropol 18: 281-292.
Richtsmeier JT, Aldridge K, de Leon VB, Panchal J, Kane
AA, Marsh JL, Yan P, Cole TM. 2006. Phenotypic
integration of neurocranium and brain. J Exp Zool
(Mol Dev Evol) 306B: 360-378.
Sergi S. 1934. Ossicini fontanellari della regione dellambda nel cranio di Saccopastore e nei crani
neandertaliani. Riv Antropol 30: 101-112.

Figure 1. Neanderthals display a relative shortening of theupper parietal profile, as possible allometric consequence
associated with encephalization and structural neurocranialconstraints. The presence of hypostotic traits at the parietooccipital boundaries may suggest morphogenetic problems
associated with a derived brain size and a plesiomorphic
braincase organization.

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

Bruner E, Manzi G, Arsuaga JL. 2003. Encephalization
and allometric trajectories in the genus Homo:
Evidence from the Neandertal and modern lineages.
Proc Natl Acad Sci USA 100: 15335-15340.
Bruner E, de la Cuétara JM, Holloway R. 2011a. A
bivariate approach to the variation of the parietal
curvature in the genus Homo. Anat Rec 294: 15481556.
Bruner E, Mantini S, Musso F, de la Cuétara JM, Ripani
M, Sherkat S. 2011b. The evolution of the meningeal
vascular system in the human genus: from brain shape
to thermoregulation. Am J Hum Biol 23: 35-43.
Gunz P, Harvati K. 2007. The Neanderthal "chignon":variation, integration, and homology. J Hum Evol 52:
262-274.
Gunz P, Neubauer S, Maureille B, Hublin JJ. 2010. Braindevelopment after birth differs between Neanderthals
and modern humans. Curr Biol 20: R921-922.

069

Virtual Endocast of Qafzeh 9, A Representative of
Early Modern Humans

Osamu Kondo1, Daisuke Kubo1, and Naomichi Ogihara2
1Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Japan

2Department of Mechanical Engineering, Faculty of Science and Technology, Keio University, Japan

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

070

Paleoneurology is an important field in the studyof 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 links of the hard
evidence to behavioral and cognitive differencesamong fossil species, which might have drove thespeciation between the taxa. In this context, C01
team of the project RNMH (Replacement of
Neanderthals by Modern Humans) aims to present
reasonable ways for reconstruction of the fossil
hominids in order to estimate the endocasts and thebrains, and also aims to make possible inferences
for behavioral and cognitive differences between
Neanderthals and modern humans, based on theestimated endocasts and brains.
In this paper, we make an endocast from thefossil cranium of Qafzeh 9 and compare the sizeand shape of it with those of modern humans
and several fossil hominids. At the same time,with assessment of the degree of deformation of
the original fossil cranium, we try to correct theobserved degree of deformation or distortion.
Qafzeh 9 specimen is a fossil representative of earlymodern Homo sapiens and a key specimen in studies
of expansion of Homo sapiens and of extinction of
Homo neanderthalensis. However, the endocast of
Qafzeh 9 was never produced.
Qafzeh 9 was discovered and excavated by B.
Vandermeersch in 1969 from the Qafzeh Cave inIsrael. It was associated with Middle Paleolithic

assemblages of the Levalloiso-Mousterian culture( Vandermeersch, 1981). 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 evenyounger classic Neanderthals. According to his
first description, the Qafzeh 9 individual is an
adult female, whose cranial bones were crashed andheavily fragmented, but the whole cranium waswell restored and reconstructed (Vandermeersch,1981). The endocranial feature was only describedon the parietal meningeal vessels because no
endocasts have been made for this specimen. The
degree of distortion was unknown, though thepresent reconstruction exhibits possible distortions
both in the neurocranium and the face (Figure1). By contrast, the endocranial capacity wascalculated to be from 1508 to 1554 cc using several
regression equations of the linear measurements(Vandermeersch, 1981), and the mean value of 1531
cc was reproduced in a textbook (Holloway et al.,2004).
We conducted a virtual reconstruction of theendocast from the CT-scan images of the specimen,with their planar resolution of 0.41 mm/pixel andthe slice thickness of 0.5 mm. The process followedthat of Kubo and Kono (2011), i.e. after extracting
the cranial model from the CT images by using athreshold between the bone and air and making aphysical mold of the endocranium with 3D printer,
we filled the gap and repaired the lacking portions

of the inner cranial surface with clay (Figure 2). The
more complete surface model of the endocast was
produced from re-scanning images of the repaired
mold. A preliminary estimation of the endocranial
volume (ECV ) was obtained to be 1411 cc. We
should proceed to comparison of the ECV and theendocast morphology with those of comparative
samples as well as assessment of the distortion andits correction.
In the preliminary study on the Qafzeh 9 cranium,
we observed the significant distortion, which was
determined as the degree of right-left asymmetry;the posterior neurocranium skews with the rightanterior and the left-posterior warping, and the facedeviates toward the right as a whole. The correction
value for the present Qafzeh 9 cranium can begiven in terms of normal RL asymmetry observedin anatomical collections of modern human skulls,
which will provide a meaningful deviation of thepresent Qafzeh 9 endocast, and may change theimplication and understanding of its endocranial
morphology.
References
Holloway RL, Broadfield DC and Yuan MS (2004) The
Human Fossil Record. vol. 3. Brain Endocasts - The
Paleoneurological Evidence. Wiley-Liss.
Kubo D and Kono RT (2011) Endocranial restorationand volume estimation of the Minatogawa IV
cranium using micro-CT and 3D printer systems.
Anthropol. Sci. 119: 203-209.
Vandermeersch B (1981) Les Hommes Fossiles deQafzeh (Israel). Editions du CNRS, Paris.

Figure 2. Physical mold of Qafzeh 9 endocranium with infilled clay.

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

Figure 1. Qafzeh 9 cranium and endocast model.

071

Cerebral Sulci and Gyri Observed on Macaque
Endocasts
Yasushi Kobayashi1, Toshiyasu Matsui1, Yoshinori Haizuka2, Naomichi
Ogihara3, Naoki Hirai4, and George Matsumura2
1Department of Anatomy and Neurobiology, National Defense Medical College, Japan
2Department of Anatomy, Kyorin University School of Medicine, Japan
3Department of Mechanical Engineering, Keio University, Japan

4Department of Integrative Physiology, Kyorin University School of Medicine, Japan

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

072

Information on the cortical subdivisions of
Neanderthal brains is essential in understanding
the cognitive capabilities of this extinct hominid.
To address this problem, we investigatedcharacteristics of skulls that reliably indicate theborders of cortical areas. In the present study, weevaluated the correlation between the impressions
on the inner surface of the skull and the cerebral
gyri and sulci.
We used 5 aged monkeys (Macaca fascicularis,
3 males and 2 females) that had long been used
for studies on the gastrointestinal tract and werescheduled to be euthanized. Under deep anesthesia
with ketamine, xylazine and sodium pentobarbital,
the animals were perfused transcardially with 2.5%
formalin in phosphate buffer (pH 7.4) for 2 min(200 mL/min), followed by 10% formalin in thesame buffer solution for 50 min (200 mL/min for
initial 10 min, followed by 100 mL/min for theremaining 40 min). The first 2 cervical vertebrae
were laminectomized, and the spinal cords were cut
at C2-C3 junction, and the heads were separated
from the neck approximately at the same leveland stored in phosphate-buffered 2.5% formalinsolution at room temperature. The heads werethen scanned using Toshiba Asterion CT scannerat the Laboratory of Evolutionary Biomechanics,
Department of Mechanical Engineering, Keio
University to obtain 0.5 mm scan slices of 512 x
512 pixels.
The skulls and brains were next analyzed
macroscopically and histologically at the National

Defense Medical College. Af ter removing
the scalp, fascia, and temporalis muscles, we
mounted the heads on a stereotaxic frame, andphotographically recorded the surface of thecalvaria. We then removed the calvaria and thedura mater, and recorded the surface of the brains.
CT images were analyzed using Amira 5.4 (Visage
Imaging, Berlin, Germany) on a Z620 workstation(Hewlett-Packard Japan, Tokyo, Japan).
Virtual endocasts that were created using
CT data showed marked pattern of cerebral
convolution (Figure 1). We observed impressions
corresponding to most of the major cerebral sulci:
the central, lateral, arcuate, principal, intraparietal,
lunate, superior temporal, and anterior middle
temporal sulci on the dorsolateral surface, as wellas the medial and lateral orbital sulci on the ventral
surface. The correspondence of the impressions
to the cerebral sulci was confirmed by comparing
the virtual endocasts with the brains of the same
animal.
We further analyzed the individual differences insulcal patterns of endocasts with those of the brain,
and observed that some of the differences can beinferred by the endocasts, particularly in the frontal
lobe: the medial and lateral orbital sulci (Figure 2),
the arcuate and the principal sulci.
Usually, adult skulls of both Neanderthals andmodern humans show hardly any impressions
corresponding to the cerebral sulci except for theorbitosphenoidal crest and digital impressionsof the orbital plate of the frontal bone. On theFigure 1. Reconstructed surface images of the skulls (A, D) and the virtual endocasts (B,C, E, F) of a male (M15, panels A-C;) and a female (M19, D-F) monkeys. Marked impressions
corresponding to major cerebral sulci were identified: ar arcuate sulcus, pr principal sulcus,
ce central sulcus, ip intraparietal sulcus, lunate sulcus, la lateral sulcus, st superior temporalsulcus, amt anterior middle temporal sulcus, lo/mo lateral and medial orbital sulci.

other hand, impressions possibly corresponding
to cerebral sulci were reported in some extant
hominids such as Australopithecus. Infant skulls of
Neanderthals exhibit some hints of convolutional
pattern on their endocasts. These findings indicatethat we may be able to identify cerebral sulci insmall skulls that have relatively thin connective
tissues intervening the skull and the brain.

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

Figure 2. The orbitofrontal surface of two female monkeys (M17, panels A-C, and M19, D-F).
Panels B and E illustrate the orbital sulci that were traced on the images of virtual endocasts (A,D). The lateral orbital sulcus of M17 joins the medial orbital sulcus on both sides (C), whereas
the lateral and medial orbital sulci of M19 run nearly parallel to each other (F). These individual
differences were well preserved on the endocasts.

073

Reconstruction of the Brain from Skull Fossil Using
Computational Anatomy

Takanori Kochiyama1, Hiroki C. Tanabe2, and Naomichi Ogihara3
1Primate Research Institute, Kyoto University & ATR-Promotions Brain Activity Imaging Center, Japan

2Graduate School of Environmental Studies, Nagoya University & National Institute for Physiological Sciences, Japan
3Department of Mechanical Engineering, Keio University, Japan

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

074

We investigated the presumed differences inlearning abilities between Neanderthals andmodern humans by combining evidence fromthe morphological analysis of fossil brains andthe functional mapping of modern humans’ brain
functions. Our study faced a fundamental challenge:
although we could easily collect various data about
the structures and functions of the modern humanbrain, the only available data for the Neanderthal’s
brain were skull CT images reconstructed fromfossilized brains produced by Project C01. Hence, toachieve our purpose, we had to establish a method
for extrapolating to human brain functions from skull
anatomy, which is the only data available for bothmodern humans and Neanderthals. In this study,
we tried to develop a skull-based image registration
method using the computational anatomy technique,a standard method such as functional MRI (fMRI)
and voxel based morphometry (VBM) used inneuroimaging research. The basic idea is as follows:First, the skull shape reconstructed from CT imagesof a fossil brain is spatially deformed to the modern
human skull shape segmented from MRI/CT
image. Nonlinear spatial transformation method isused to estimate a globally one-to-one smooth andcontinuous mapping between these skull shapes.
Diffeomorphic Anatomical Registration ThroughExponentiated Lie algebra or DARTEL registration
method is a promising candidate for our research.
Using the deformation field estimated by thenonlinear transformation, the modern human cerebral
cortex segmented from MRI image is then inversetransformed to construct virtual Neanderthal’s cortex

which is consequently best fitted to its skull shape.
This deformation is also applied to the functional
brain map of modern human to presume the
Neanderthal’s one. The schema of this study is shownin Figure 1.
To validate our proposed strategy, we currently
conduct a simulation analysis using modern human
MRI. In this report, we compared the accuracy ofthe skull-based brain exchange between two modern
human individuals with the brain (gray matter; GMand white matter; WM) -based exchanging. The
procedures are as follows: Fifteen healthy subjects
participated in this study. All MRI images wereacquired using a 3-Tesla MR scanner (Siemens
Allegra) at the National Institute for Physiological
Sciences. A T1-weighted fine structural wholebrain image was acquired using a magnetization
prepared rapid acquisition gradient echo (MPRAGE). The parameters were as follows: TR=2500
ms; TE=4.38 ms; inversion time = 1100ms; FA = 8°;
FoV = 230 mm; number of slices per slab = 192; voxel
dimensions = 0.9 × 0.9 × 1.0 mm. Image processing
was performed using the Statistical Parametric
Mapping package (SPM8; Wellcome Departmentof Cognitive Neurology, London, UK) implemented
in MATLAB (Mathworks Inc., Sherborn, MA,
USA). First, we segmented individual MRI images
into GM (gray matter), WM (white matter) and
Skull by segmentation-normalization algorithm in
SPM. Then, using DARTEL registration method,
the subj2-to-subj1 transformation was estimatedwith matching of their skull images or brain (GMand WM) images. Finally, inverse transformation

Figure 2. Result of the brain exchange.

Figure 1. The schema of this study.

the following issues: (1) we quantitatively evaluate thesimilarity and difference in local features of the corticalsurface between individuals. Deformation based
morphometry analysis can be suitable for this purpose.
This information might help to compensate for the
lower resolution of the skull-based transformation.
(2) we expand this method to handle the spatial
registration between MRI and CT. Our goal is toestablish the method for casting a modern human’s
cerebral cortex (MRI) from MRI in a Neanderthal'sskull (CT) with matching between the modern
human’s skull (MRI or CT) and the Neanderthal'sskull (CT). Hence, we should update our method toallow the multimodal image processing. We validate
the accuracy of the method and clarify the problems inapplication to fossil brain CT.

Towards Digital Reconstruction of Fossil Crania and their Brain Morphology

from subj1 to subj2 was applied to subj1’s brain toconstruct the deformed subj1’s brain in the shape ofsubj2’s brain.
Figure 2 shows the result of the brain-based (uppercolumn) and skull-based (lower column) exchange.
The left and right row indicates the original MRI
image for subject 1 and 2, respectively. The middle
row shows the transformed subj.1 image, with theshape of subj2’s brain. In the case of the brainbased matching, not only global features but also
the local sulcus and gyrus arrangements were wellmatched between the transformed (upper middle)
and the original subj2’s image (upper right). In thecase of the skull-based matching, the global skull
shape and the cortical surface shape were similar
between the transformed image and the original
image. However, there was obvious misalignment
in the local structures as well as deep structures (e.g.
size and shape of ventricle). To access the average
accuracy of the transformation, we repeatedly applied
this analysis to the pair of subject 2 and one of theremaining 14 subjects and then averaged the virtual
subj2’s brain images. We found that the global shapeof the brain was well converged on that of subj2’s
brain even after skull-based matching, however, interms of local accuracy, the only prominent landmark
structures such as the lateral and central sulci could
be identified on the averaged brain.
Following the results of this simulation and taking
into account the application of this method to CTimage of a fossil brain, we will further investigate

075

Innovative Society and Social Brains

Hiroki C. Tanabe

Graduate School of Environmental Studies, Nagoya University, & National Institute for Physiological Sciences, Japan

Neuroimaging Approach to the Learning Hypothesis

076

Research team C02 adopts cognitive neuroscientificand comparative neuroanatomical approaches to
examine our hypothesis that the replacement of
Neanderthals by Homo sapiens was promoted by
the differences in their learning abilities (‘learning
hypothesis’). By assuming that morphologicalchanges in fossil skulls reflect functional differences
between modern human brains and Neanderthal’s
brains, we try to identify gaps in learning abilities
based on differences in cerebral morphology andregion-specific activities using our developed analysis
scheme (i.e. integrated system of computational
neuroanatomy and brain function mapping).
From early 1990’s, advances in magnetic
resonance imaging (MRI) techniques havefacilitated functional brain imaging and direct
measurement of living human brain activities (i.e.
functional MRI). Such techniques have enabled
us to map the specific functions of the modern
human brain. One of our team’s goal is to create
the maps regarding the difference of ‘learning
abilities’ between Neanderthals and Homo sapiens.
According to Tomasello (1999), some individual
or group of individuals first invented a primitive
version of the artifact or practice (i.e. creative or
innovative activity; what we call individual learning
ability), and then some later user or users madea modification or adopted without change formany generations (i.e. learned and used by others
over time; what we call social learning ability) in
Homo sapiens history. The important point is thatthe Homo sapiens is able to pool their cognitive
resources, and the base of this ability is that they

can understand conspecifics as beings like themselves
who have intentional and mental lives like their own(Tomasello et al., 1993). In line with this argue, we
assume that innovative society is deeply correlatedwith the two components of the cognitive ability:‘intrinsic drive (internal motivation and perspective)
to produce creative activity’ and ‘social cognition andinteraction’.
In the present talk, I focus on the neural substratesof ‘social brains’. Social brain is a circumscribed
set of brain regions that are dedicated to the social
cognition and interaction (Brothers, 1990). The
most important attribute of the social brain is tomake predictions about other’s actions on the basisof their mental states, and this ability is developed
from early stage of human life. As an exampleof these social cognitive abilities, I introduce ourneuroimaging studies those investigate neuralsubstrates of ‘eye contact’ and ‘joint attention’ (Saito
et al., 2010; Tanabe et al., 2012). Both of them
are the merkmals of the social cognitive abilities
during the early development of humans. Eyecontact affords to establish a communicative link
between humans, and prompts joint attention.
Joint attention is an ability to coordinate attention
between interactive two persons regarding objects
or events. The impaired development of it is a
cardinal feature of autism spectrum disorder (ASD).
As the eye contact is implicated in the sharing ofvarious psychological states, it might provide a
communicative context in which joint attention canbe initiated. To elucidate the neural mechanisms ofinter-subjective sharing during eye contact and jointbetween Homo sapiens and Neanderthals.

References:
Brothers L (1990) The social brain: A project for
integrating primate behaviour and neurophysiology ina new domain. Concepts Neurosci., 1, 27-251.
Saito DN, Tanabe HC et al. (2010) “Stay tuned”: Interindividual neural synchronization during mutual gazeand joint attention. Front. Integ. Neurosci., 4, 127.
Tanabe HC, Kosaka H et al. (2012) Hard to “tune in”:
Neural mechanisms of live face-to-face interaction

with high-functioning autism spectrum disorder.
Front. Hum. Neurosci., 6, 268.
Tomasello, M. (1999) The Cultural Origins of Human
Cognition. Harvard Univ. Press.
Tomasello, M., Kruger, A.C., Ratner, H.H. (1993)
Cultural learning. Behavioral and Brain Sciences, 16,495-552.

Neuroimaging Approach to the Learning Hypothesis

attention, we conducted a hyper-scanning functional
MRI experiment while they were engaged in joint
attention tasks with eye contact as the baseline. We
assume that the eye contact based psychologicallyshared mental states is neurally represented by theinter-subject synchronization of the ‘state’ of thebrain activity, which is obtained by the eliminationof the task-related activation component. In
contrast, we detected brain regions related tojoint attention in terms of task-related activity.
Moreover, in order to compare the difference ofneural mechanisms during eye contact and joint
attention between typically-developed young
adult individuals and those with high-functioning
ASD, we try to elucidate brain regions those play
important role in social cognition during eye-based
communication. Our findings suggest that the rightinferior frontal gyrus (IFG) plays an importantrole for shared intention during eye contact thatprovides the context for joint attention. Taken withother neuroimaging studies, we think that the right
IFG is one of specific regions regarding the social
cognition, and is a candidate for the difference077

Social Neuroscience of Stone Tool-Making Skill
Acquisition

Dietrich Stout
Emory University, USA

Although there is a burgeoning neuroscience oftool use, there is little that might be properly calleda neuroscience of technology. Anthropological andarchaeological approaches to technology can help
to provide a broader theoretical and evolutionary
context. “Technology” is a fuzzy category thatloosely corresponds to human actions that 1)
involve the use or modification of objects, 2)
display a complexly organized multi‐level structure,and 3) are socially reproduced. These characteristics
may be better understood with reference toneuroscience research on perceptual‐motor control,

Neuroimaging Approach to the Learning Hypothesis

078

object manipulation, motor resonance, imitation
learning and goal‐directed action. Research using
neuroscience methods (PET, fMRI) to examine
the archaeologically visible behavior of stonetool‐making has provided insights into the pastrole of technology in human cognitive and brain
evolution, including relations with the humanfaculty for language. A key focus of this researchis to investigate mechanisms for the observational
understanding and social reproduction of stonetool‐making skills.

Study Implication Related to Sense of Acceptance:
Key Factor of Social Learning
Hiroaki Kawamichi1, Kazufumi Yoshihara1, Ryo Kitada1, 2, Masahiro Matsunaga1,
Akihiro Sasaki1, Yumiko Yoshida1, 2, Haruka Takahashi,1, 2 and Norihiro Sadato1, 21National Institute for Physiological Sciences, Japan 2The Graduate University for Advanced Studies

Recent advances of neuroimaging techniques,
such as functional magnetic resonance imaging(fMRI), have revealed the neural correlates
underlying a wide range of social abilities including
a sense of acceptance. In this paper, we will
review research related to the sense of acceptance,
including the modulation of brain activation.
Finally, we will introduce our future experimental
plan.
Sense of acceptanceSociometer theory has proposed that social
acceptance from other people enhances selfesteem (Leary et al., 1995), and self-esteem is aconsequence of the degree to which people receive
acceptance from other people. As proposed inperson-centered therapy, a sense of acceptance can
influence on social behaviors through psychologicalcontact (Rogers, 1957). Congruence, unconditional
positive regard, and an empathic attitude of thetherapist are the most important factors in personcentered therapy. Within successful therapy, clientchanged his / her way of thinking. This change ofway of thinking promotes new social behaviors.
Thus, perceiving a sense of acceptance contributes
promoting new social behaviors through changing
a way of thinking.
Functional MRI studies have shown that
acceptance (praise) from others activates thestriatum, which is part of the reward system (Izuma
et al., 2008). On the other hand, rejection fromothers shows activation in the dorsal anterior
cingulate cortex (ACC), part of the pain matrix

Neuroimaging Approach to the Learning Hypothesis

Introduction
Cognitive and behavioral capabilities of modern
humans, such as creativity, likely played a key rolein the demise of Neanderthals. Both individual
and social learning contribute to this creativity. For
instance, in regards to individual learning, humans
acquire new skills through trial and error processes;
thus, this learning ability facilitates the inventionof new skills and technologies. In social learning,on the other hand, humans acquire new skills byobserving (Bandura, 1965) and imitating (Field& Walden, 1982) the behaviors of others. Social
learning ability thus promotes the spread of the
new skills and technologies invented by individual
learning. Predominance of these two abilities ofmodern humans in comparison with Neanderthals
might lead to replacement of Neanderthals bymodern humans.
Modern humans have enhanced social abilities
compared with other animal species (Dunber,1998); indeed, the capacity for social learning
is one of the major characteristics of modern
humans. There are four necessary conditions ofsocial learning processes: paying attention to atarget, the retention and reproduction of a target’s
behavior, and motivation for imitation (Bandura,
1977). Acceptance towards others’ behaviors
contributes to the attention to the retention andreproduction the behavior. In addition, acceptancefrom others, i.e., approval, enhances the motivation
to imitate the behaviors (Bandura, 1977). Thus,
sensing acceptance from others might facilitatesocial learning processes.

079

(Eisenberger et al., 2003), and is accompanied
by a self-esteem decrease (Eisenberger et al.,2011a). Interestingly, social support has beenshown to decrease psychological pain-related
activation in the ACC during social exclusion(Onoda et al., 2009). Thus, social acceptance fromothers modulates brain activation in two ways:
enhancement for reward-related activation andattenuation for pain-related activation.
Humans can also perceive a sense of acceptanceeven without the explicit words of others (Rogers,
1957), which likely stems from their nonverbal
attitude. Perceived social support can come in
the form of interpersonal touch (Coan et al.,2006). For instance, hand-holding with a spouseor stranger decreases the neural response to pain
threat in the right anterior insula, superior frontal
gyrus, and hypothalamus, including the affective
pain matrix (Coan et al., 2006). In addition, aphoto of romantic partner also decreases painrelated responses (Eisenberger et al., 2011b).
Therefore, a sense of acceptance stemming fromthe nonverbal and verbal behaviors of others canmodulate emotional related responses during social
behaviors.

Neuroimaging Approach to the Learning Hypothesis

080

Suggested experiment on the sense of acceptanceRemained issue:
Despite the progress in research on the sense of
acceptance, there are still issues that need to beresolved. First, as mentioned in the introduction,a sense of acceptance enhances social behavior.
However, the neural correlates underlying thelink between a sense of acceptance and theenhancement of social behavior is unresolved.
Second, the motives for social behavior mainly
consist of psychological factors. Thus, the link
between negative feelings stemming frompsychological factors and a sense of acceptancewarrants further investigation.
Example of experimental design outline:
I n t h i s a r t i c l e, we s u g ge s t a p o t e n t i a lexperimental design regarding the link between
negative feelings stemming from psychologicalfactors and a sense of acceptance. In this type ofexperiment, empathic pain, the negative feelingaroused by seeing others in pain (Singer et al.,2004), can be treated as psychological pain. Other
conditions in the experiment include supportive
words or hand holding as a sense of acceptance.
For this experiment, each subject observes painful
or non-painful visual stimuli throughout thesession. They then evaluate the stimuli while theleft hand of a familiar person is placed on theirhand (human hand condition). Alternatively, thecontrol condition is when a non-human (e.g.,rubber) hand or no hand (non-hand condition) isused.

Figure 1. Time chart example of visual stimuli
presentation.

Conclusion
In this paper, we have indicated the importanceof a sense of acceptance for promoting social
learning. The sense of acceptance has been
investigated in wide range of studies including its
underlying neural mechanisms in regards to social
behavior or psychological factor effects. However,there is a missing link at the mechanistic level
among social behavior, psychological factors, andsense of acceptance. Our proposed experiment may
help clarify the neural mechanisms underlying
social learning.
References
Coan, J.A., Schaefer, H.S., & Davidson, R.J. (2006)
Lending a hand: social regulation of the neural
response to threat. Psychological Science, 17, 10321039.

reduced social pain during emotional support. Social
Neuroscience, 4, 443-454

Neuroimaging Approach to the Learning Hypothesis

Decety, J. (2011) Dissecting the neural mechanisms
mediating empathy. Emotion Review, 3, 92-108.
de Waal, F.B.M. (2008) Putting the altruism back into
altruism: the evolution of empathy. Annual Review
of Psychology, 59, 279-300.
Dunbar, R.I.M. (1998) The Social Brain Hypothesis.
Evolutionary Anthropology, 6(5), 178-190.
Eisenberger, N.I., Inagaki, T.K., Muscatell, K.A., Haltom,
K.E.B., & Leary, M.R. (2011a) The neural sociometer:
brain mechanisms underlying state self-esteem. Journalof Cognitive Neuroscience, 23, 3448-3455.
Eisenberger, N.I., Lieberman, M.D., & Williams, K.D.
(2003) Does rejection hurt? An fMRI study of social
exclusion. Science, 302, 290-292.
Eisenberger, N.I., Master, S.L., Inagaki, T.K., Taylor, S.E.,
Shirinyan, D., Lieberman, M.D., & Naliboff, B.D.
(2011b) Attachment figures activate a safety signalrelated neural region and reduce pain experience.
Proceedings of National Academy of Sciences USA,108, 11721-11726.
Field, T.M. & Walden T.A. (1982) Production andperception of facial expressions in infancy and earlychildhood. Advances in Child Development andBehavior, 16, 169-211.
Izuma, K., Saito, D.N., & Sadato, N. (2008) Processingof social and monetary rewards in the humanstriatum. Neuron, 58, 284-294.
Leary, M.R., Tambor, E.S., Terdal, S.K., & Downs, D.L.
(1995) Self-esteem as an interpersonal monitor: thesociometer hypothesis. Journal of Personality andSocial Psychology, 68, 518-530.
Kuhn, S., Muller, B.C., van Baaren, R.B., Wietzker, A.,
Dijksterhuis, A., & Brass, M. (2010) Why do I likeyou when you behave like me? Neural mechanisms
mediating positive consequences of observing
someone being imitated. Social Neuroscience, 5,384-92.
Preston, S.D. & de Waal F.B.M. (2002) Empathy: its
ultimate and proximate bases. Behavioral and Brain
Sciences, 25, 1-20.
Rogers, C.R. (1957) The necessary and sufficientconditions of therapeutic personality change. Journalof Consulting Psychology, 21, 95-103.
Singer, T., Seymour, B., O’Doherty, J., Kaube, H.,
Dolan, R.J., & Frith, C.D. (2004) Empathy for pain
involves the affective but not sensory components ofpain. Science, 303, 1157-1162.
Onoda, K., Okamoto, Y., Nakashima, K., Nittono, H.,
Ura, M., & Yamawaki, S. (2009) Decreased ventral
anterior cingulate cortex activity is associated with081

Neural Substrates of Motivation to Learn and Fatigue

Kei Mizuno

RIKEN Center for Molecular Imaging Science, Japan

Neuroimaging Approach to the Learning Hypothesis

082

Differences in learning abilities are thought
to be demonstrated by elucidating the brain
regions involved in individual and social learningof modern humans using neuropsychologicalmethods, and by comparing the differencesof brain regions between modern and ancienthumans based on fossil reconstruction of ancienthuman brains. The study project for Replacementof Neanderthals by Modern Humans (RNMH)
has focused on the neural bases of learning control(enforced learning) in individual learning and of
imitative learning in social learning.
Although motivation to learn is an enforced
factor for individual and social learning, the neuralsubstrates associated with motivation to learn havenot previously been determined. Therefore, weinvestigated the neural substrates of motivation tolearn in modern humans using functional magnetic
resonance imaging (fMRI). We confirmed thatsense of accomplishment was an intrinsic reward
is associated with motivation to learn (Yoshida etal, 2008). In addition, the striatum was strongly
activated by sense of accomplishment, and striatal
activation levels were positively associated witha motivation-to-learn score derived from aquestionnaire (Mizuno et al., 2008). As academic
rewards for motivational arousal, there are sensesof accomplishment and competence, self-efficacy,
and acceptance by others (Deci et al., 1991, 1996).
Neural bases of acceptance by others have been
also reported using fMRI. Perception of a good
reputation by others induces activations of themedial prefrontal cortex and striatum (Izuma etal., 2008). These results suggest that striatum plays
crucial role for motivation to learn.
In children and adolescents, there is a negative
correlation between motivation to learn andfatigue, which is likely to be two sides of thesame coin. This finding indicates that fatigue isa negative reinforce to motivation to learn. Our
collaborators revealed that the medial orbitofrontal
cortex was associated with fatigue sensation
using positron emission tomography (Tajima
et al., 2010). Recently, we also confirmed that
activity of the medial orbitofrontal cortex wasincreased depending on time on fatigue-inducing
task using fMRI. In addition, the activation ofmedial orbitofrontal cortex was attenuated by theperception of intrinsic reward, suggesting that
intrinsic reward modulates fatigue alleviation level.
In addition to clarify the neural bases associatedwith motivation to learn and fatigue, we tried todetermine what cognitive function is correlatedwith level of motivation to learn and/or fatigue.
The neural substrates of that cognitive function
may provide evidence for discussion of thedifferences in learning abilities between modern
and ancient humans based on the differences inthe brain morphologies and functions between
them. In children and adolescents, we explored theassociation between motivation to learn and othercognitive functions, and found that motivation tolearn was positively correlated with performanceon a divided attention task (Mizuno et al., 2011a).
We used kana pick-out test (Kaneko, 1996), which
is one of the verbal dual task, to evaluate the ability

References
Deci, E. L., R. J. Vallerand, L. G. Pelletier and R. M.
Ryan 1991 Motivation in education: the selfdetermination perspective. Educational Psychologist
26(3-4): 325-346.
Deci, E. L., R. M. Ryan and G. C. Williams 1996 Needsatisfaction and the self-regulation of learning.
Learning and Individual Differences 8(3): 165-183.
Izuma, K., D. N. Saito and N. Sadato 2008 Processing ofsocial and monetary rewards in the human striatum.

Neuron 58(2): 284-294.
Kaneko, M. 1996 A neuropsychological index of aging
data from 9000 elderly people. Nippon Ronen Igakkai
Zasshi 33(11): 811-815. (in Japanese)
Mizuno, K., M. Tanaka, A. Ishii, H. C. Tanabe, H. Onoe,
N. Sadato and Y. Watanabe 2008 The neural basisof academic achievement motivation. NeuroImage
42(1): 369-378.
Mizuno, K., M. Tanaka, S. Fukuda, K. Imai Matsumura
and Y. Watanabe 2011a Relationship between
cognitive function and prevalence of decrease
in intrinsic academic motivation in adolescents.
Behavioral and Brain Functions 7: 4.
Mizuno, K., M. Tanaka, S. Fukuda, K. Imai Matsumura
and Y. Watanabe 2011b Relationship between
cognitive functions and prevalence of fatigue inelementary and junior high school students.Brain &Development 33(6): 470-479.
Mizuno, K., M. Tanaka, S. Fukuda, T. Sasabe, K. ImaiMatsumura and Y. Watanabe 2011c Changes incognitive functions of students in the transitional
period from elementary school to junior high school.
Brain & Development 33(5): 412-420.
Mizuno, K., M. Tanaka, H. C. Tanabe, N. Sadato and Y.
Watanabe 2012 The neural substrates associated withattentional resources and difficulty of concurrent
processing of the two verbal tasks. Neuropsychologia
50(8): 1998-2009.
Tajima, S., S. Yamamoto, M. Tanaka, Y. Kataoka, M.
Iwase, E. Yoshikawa, H. Okada, H. Onoe, H.
Tsukada, H. Kuratsune, Y. Ouchi and Y. Watanabe
2010 Medial orbitofrontal cortex is associated with
fatigue sensation. Neurology Research International
2010: Article ID 671421.
Yoshida, M., M. Tanaka, K. Mizuno, A. Ishii, K. Nozaki,
A. Urakawa, Y. Cho, Y. Kataoka and Y. Watanabe
2008 Factors influencing the academic motivation ofindividual college students. International Journal of
Neuroscience 118(10): 1400-1411

Neuroimaging Approach to the Learning Hypothesis

to divide attention. In contrast to the motivation
to learn, fatigue was negatively correlates with theperformance of divided attention task (Mizuno
et al., 2011b). In addition, we found that ability
to divide attention was dramatically developed
from childhood to adolescence (Mizuno et al.,2011c). Taken together, these results suggest that
motivation to learn and fatigue influence the agedependent development of cognitive function inchildren and adolescents.
The fMRI study revealed that processing ofdivided attention acquires activation of the left
dorsal inferior frontal gyrus and superior parietal
lobule in young adults (Mizuno et al., 2012). In
children and adolescents, we also found that thesebrain regions were activated during the same
divided attention task, and the activation levels areaffected by the extent of motivation to the task andbaseline fatigue. These results suggest that increased
activations of these frontal and parietal regions
during dual task performance may be associatedwith the capacity for attentional resources. In
modern humans, it is known that these cortices
develop from childhood to adulthood.
These findings highlighted the regions of interestof the brain, such as striatum and orbitofrontal,
frontal and parietal cortices, in relation to
motivation to learn, fatigue and cognitive
function affected by motivation and fatigue inmodern humans. Morphological and functional
comparisons of these regions between modern
and ancient humans are expected to help elucidate
differences in learning abilities between modern
and ancient humans. We expect these outcomes tocontribute the achievement of the RNMH project.

083

P o s t e r
Poster Presentations

Group A01: Palaeolithic Archaeology						Group A02: Hunter-Gatherer Studies						
Group B01: Evolutionary Models							Group B02: Climate and Geoscience							Group C01: Skull and Brain								Group C02: Neuroscience

RNMH2012

Aimed Spear-Throwing in Modern Humans: A
Preliminary Study of Upper Limb Kinematics

Yasuo Higurashi

Graduate School of Human Sciences, Osaka University, Japan

Palaeolithic Archaeology Group A01

086

Introduction
Humans have the ability to throw with speed andaccuracy. A few non-human primates, such as tufted
capuchins (Westergaard et al., 1994), Japanese macaques(Leca et al., 2008), and chimpanzees (Osvath, 2009),
reportedly also have the ability to throw. Although Isaac(1987) stated that throwing behavior in the context ofhuman evolution had drawn little scholarly attention,
this situation is changing. Recently, biomechanicalmethods have been used to identify anatomical factorsthat affect the throwing performance of modern
humans and to evaluate the throwing ability of earlier
Homo species (Roach et al., 2012).
The existence of anatomical features that enhancethrowing performance does not necessarily mean thatthe species used throwing-based projectile weapons inhunting. The spear thrower (atlatl or woomera), whichis unequivocal evidence of hunting with a projectile
weapon, came into use after the disappearance of Homo
species other than modern humans (see Schmitt et al.,2003 and references therein). The oldest known spears
were discovered in Schöningen, Germany, and dated
to 400,000 years ago, the time of Homo heidelbergensis
or the early Neanderthals in Europe (Thieme, 1997;
for evolutionary scenarios for the origin of the
Neanderthals, see Rightmire, 2008 and Hublin, 2009).
Whether these spears were throwing- (Thieme, 1997;
Rieder, 2003) or thrusting-based weapons (Schmitt etal., 2003) is controversial.
I am currently undertaking a research project tobetter understand throwing behavior, particularly spear
throwing, in Neanderthals and modern humans. In thispresentation, I report the results of two experiments:
electromyography (EMG) of the biceps brachii during

forearm supination, and upper-limb kinematics in spear
throwing with a spear thrower.
Experiment 1: EMG of the biceps brachii during
forearm supination
There are a number of features in which themorphology of the Neanderthal postcranial skeleton
differs from that of modern humans (Trinkaus, 1983;
Aiello and Dean, 2002; Weaver, 2009). Some of thesefeatures are unique to the Neanderthals (autapomorphic),
whereas others are shared with earlier hominins(symplesiomorphic) (Weaver, 2009).
In the Neanderthals, the radius had a more medially
positioned radial tuberosity than that found in modern
humans (Trinkaus and Churchill, 1988; De Groote,2011). A medially positioned radial tuberosity is also
observed in the apes, particularly in the orangutans
and chimpanzees (Aiello and Dean, 2002). The radial
tuberosity provides the insertion for the biceps brachii
muscle, which serves as a supinator of the forearm as
well as a flexor of the elbow. However, an EMG study
carried out by Travill and Basmajian (1961) showedthat this muscle generally remains inactive duringslow supination and during supination with the elbow
extended. The medially positioned radial tuberosity in
the Neanderthals may have maintained the momentarm of the biceps brachii for supination throughout
forearm rotation, suggesting that the Neanderthals had
more power in pronation with the elbow flexed than domodern humans (Trinkaus and Churchill, 1988).
Here, I studied the activity patterns of the biceps
brachii during forearm supination in more detail. Many
parts of this experimental protocol were adapted from
Travill and Basmajian (1961). The EMG activity wasFigure 1. In Experiment 1, a participant performed a task whilelying supine with the shoulder abducted at 90°. The elbow was
flexed at 0° (no flexion), 30°, 60°, 90°, and 120°. The amount ofelbow flexion is illustrated in gray.

Figure 2. Raw EMG traces of the biceps brachii during fast
forearm rotation in a participant. Pronation and supination
were alternately performed as fast as possible at 1-second
intervals. Gray vertical lines roughly correspond to the onset ofsupination.

Experiment 2: Upper-limb kinematics in spear
throwing with a spear thrower
I studied body movements during spear throwing
with a spear thrower to explore the following topics:
mechanics of the spear thrower
works, upper-limb kinematics,
and anatomical differences
between Neanderthals andmodern humans that might
affect their throwing ability.
Participants were instructedto throw a spear-like object at
a target using a spear thrower(F igures 3 and 4). V ideo
recordings were made using
digital camcorders. To aid indata analysis, spherical reflective
markers were placed on bonylandmarks of the body, and the
movement of these markers wasrecorded using motion-analysis Figure 3. A spear thrower
from the top view (left)
software and digitized.
and the side view (right).

Figure 4. A spear thrower in use.

Palaeolithic Archaeology Group A01

recorded at 1 kHz from the dominant biceps brachii
using bipolar surface electrodes. Simultaneously with
EMG measurements, video recordings were made at
60 Hz using a digital camcorder. While performing
forearm rotation tasks, the participant lay supine on apadded table and abducted the dominant shoulder at
90° (Figure 1). Each task involved 20 repetitions of aseries of movements that started with the forearm ina full supination, moved to full pronation, and thenreturned to the start position (supination). Forearm
rotation tasks were carried out at two speeds (slow andfast) with the elbow flexed at five different angles (0°,30°, 60°, 90°, and 120°). Under the slow-speed condition,
the participant performed every forearm pronation orsupination for 1 second. Under the fast-speed condition,
the participant performed these movements as fast as
possible at 1-second intervals.
Generally, the biceps brachii muscle was active during
supination and remained inactive during pronation.
During supination at the slow speed, the biceps brachii
was active when the elbow was flexed at 60° to 120°,
but the amplitude of the activity was very small. During
supination at the fast speed, the biceps brachii wasactive with the elbow flexed at any angle (Figure 2).
One-way ANOVA showed that the maximum EMG
activity at the fast speed was significantly higher at 120°
than at any other angle of elbow flexion (all values, p <0.05). Although the difference did not reach statisticalsignificance, the maximum activity was higher at 90°
than at smaller angles of elbow flexion.
This experiment generated both similar and different
results compared with those of Travill and Basmajian(1961). The difference might be due to the small
number of participants in this experiment or the typesof electrodes used. Nevertheless, the results support theirfinding that the biceps brachii shows marked activity
during fast supination with the elbow flexed.

087

Neander DB 2.0: Improvement of the “Replacementof Neanderthals by Modern Humans” Lithic Industry
Database
Yasuhisa Kondo1,2

1Department of Computer Science, Tokyo Institute of Technology, Japan
2Japan Society for the Promotion of Science, Japan

Palaeolithic Archaeology Group A01

088

This paper reports the recent progress of the“Replacement of Neanderthals by Modern
Humans” lithic industry database (Neander DB)
project of the research team A01 (archaeology).
The aim of this project is to map and analyse thespatio-temporal processes of the replacement of
Neanderthals by anatomically modern humans(AMHs) in higher resolution than in the previous
researches. For this purpose, members of thedatabase team are collecting the information onarchaeological sites and lithic industries in Africa
and Eurasia between 200 k.a. and 20 k.a.[1] As of
September 30, 2012, Neander DB has recorded a
total of 376 lithic industries from 3,832 culturallayers of 1,977 sites and 5,182 radiometric samples.
These data was compiled from a total of 547
literature sources including excavation reports,
research papers, and monographs. In addition to
the European databases published [2], Neander
DB is covering the whole part of Africa, Asia, andNear Oceania.
An increasing number of records have led us
to realise inter-regional inconsistencies in theterminology of lithic industries, spatial unit, andperiodisation. First, lithic industries sometimes
have different local or regional terms. For
example, Mousterian is known as the industry
that belonged to Neanderthals and has dozens ofvariant names. Some of these indicate typologicalor technological variants (Denticulate Mousterian,
Levallois-Mousterian, etc.), while others predicate
regional (Altai Mousterian, Zagros Mousterian,etc.) or temporal vatiation (Late Mousterian,

Final Mousterian, etc.). Second, the concept of“cultural layers” has originally been developedin the Levant and Europe because there are anumber of Palaeolithic cave or rock-shelter sites
in which anthropogenic deposits are geologicallylayered. However, it is difficult to apply the same
concept to open sites in Siberia and other regions.
Third, different durations of period are referred indifferent regions and different fields of research.
W hen attempting to fit the archaeologicalchronology in that of the palaeoenvironmental
sciences, we should refer to MIS (marine isotope
stage) as common time scale.
In order to fix these problems, the metadata
structure of Neander DB was thoroughly reviewed
and the numerical cardinalities between the tablesof site (cultural layer), lithic industry, and period
were reorganised. In the new version, Neander
DB 2.0 (Figure 1), one cultural layer is related to
one and more lithic industries, and each industry
belongs to one and more regions, regionallydefined time periods, and MIS. This alternation
also allows us to describe multiple industries inopen sites where the distinction of geological layers
is unclear.
Neander DB 2.0 was launched in July 2012.
With clarified metadata structure, it successfully
provides more accurate text-based retrieval of theattributes such as periods, lithic industries, andpresence of fossils (Figure 2). In my view, Neander
DB 2.0 will act as a powerful analytical tool tovisualise and understand the spatio-temporal
pattern of the “replacement” of Neanderthals

by AMHs in se ver al par ts of Eur asia. By
collaborations with informatics specialists of the
project headquarters in near future, an online
database and map architecture with associative
retrieving system will be developed for the next
generation database, Neander DB 3.0, which will
be launched when we publish the dataset at the
final phase of the project.
[1] Kondo, Y., S. K adowaki, H. K ato, M.
Naganuma, A. Ono, K. Sano, and Y. Nishiaki(2012) Network computing for archaeology: a casestudy from the "Replacement of Neanderthals byModern Humans” lithic industry database project.
In: M. Zhou et al. (eds.) Revive the Past: Proceedingof the 39th Conference on Computer Applications andQuantitative Methods in Archaeology, Beijing, 1216 April 2011. Amsterdam: Amsterdam University
Press, pp. 217-226. http://dare.uva.nl/aup/nl/
record/412958[2] D'Errico, F., W. E. Banks, M. Vanhaeren, V.
Laroulandie, and M. Langlais (2011) PACEA georeferenced radiocarbon database. PaleoAnthropology
2011: 1-12. doi: 10.4207/PA.2011.ART40.

Figure 1. User interface of Neander DB 2.0 and schematic diagram of the entity relationship.

Palaeolithic Archaeology Group A01

Figure 2. Location of the paleoanthropological sites recorded in Neander DB 2.0 (as of September 1, 2012).

089

The Initial and Early Upper Paleolithic of theJapanese Archipelago

Hiroto Nakata

College of Literature, Aoyama Gakuin University, JapanIn Paleolithic studies of Japan, we eliminated all
the “fake” materials and then re-started to clarify theInitial and Early Upper Paleolithic (hereafter, IUP
and EUP respectively) or lithic industries earlier thanthe IUP and EUP. One of the current issues is how
we correctly identify lithic artifacts in consideringlithic industries preceding the Upper Paleolithic. To
identify lithics, we have to observe individual pieces
carefully, but the contexts in which lithics are foundare also important.
The earliest lithic industry in Japan has been foundin the Tachikawa loam layer X, dated to 35ka, inthe Kanto district. In this industry, I identified three

Palaeolithic Archaeology Group A01

090

lithic technologies: heavy duty tool technology, flake
technology and blade technology. Then, I sub-divided
each of these technologies according to retouched
tool types and further technological features.
Figures 1 and 2 show that the IUP of Japan
encompassed a variety of lithic technologies, among
which notable are assemblages with denticulates
and ventrally retouched scrapers as well as thosewith a block core reduction technology. Denticulates
are associated with ventrally retouched scrapers inKyusyu. Because these assemblages do not include
trapezoids, backed blades, or blade technology, they
are likely to precede in date the Upper Paleolithic

Figure 1. Major lithic technologies of the Japanese Upper Paleolithic.

in the Kanto district. The block core technology issimilar to the “chert” flake industry found in the
Tachikawa loam layer X in Kanto. These industries
with denticulates/ventrally retouched scrapers
or block core technology do not appear to havecontinued in the Kanto district during the EUP.
They were replaced with assemblages with trapezoids,
backed blades, produced by flake technology or blade
technology, and edge-ground tools. On the other
hand, the denticulate and scraper industry continued
in Kyusyu.
Presently, it is difficult to determine whether thesetechnological differences between Kanto and Kyusyu

represent social traditions or adaptations that differed
between the two regions. For now, we stress that
there was a large variety of lithic industries in Japan
during the IUP. This implies that the replacementof the old technology, characterized by denticulates
and scrapers, with the following blade technology in
the IUP occurred over a period of their co-existence.
However, no biological evidence is currently available
regarding the producers of these lithic industries.
I’m grateful to Dr. Seiji Kadowaki and Mr. Takehiro
Miki for their assistance in preparing the English
manuscript. I’m, however, responsible for the contentsof the paper.

Palaeolithic Archaeology Group A01

Figure 2. Chronological occurrences of selected technological and behavioral elements in the Japanese Upper
Paleolithic.

091

The Archaeological Database for the RNMH Project

Yoshihiro Nishiaki

The University Museum, The University of Tokyo, Japan

Palaeolithic Archaeology Group A01

092

A team of archaeologists working on the
RNMH project has been constructing an extensive
archaeological database (called NeanderDB) to
determine temporal changes in the distribution of
Neanderthal and early modern human sites on a
global level. The database is intended to facilitate
discussion on time-space contexts and the natureof the replacement process of the Neanderthals
by early modern humans (Kondo et al., 2012;
Nishiaki, 2012). The team consists of the following
members and their collaborators:
Director: Yoshihiro Nishiaki (The University ofTokyo)
Programming: Yasuhisa Kondo (Tokyo Institute ofTechnology)
Africa and West Asia: Seiji Kadowaki (Nagoya
University)
Europe: Katsuhiro Sano (Tohoku University) andAkira Ono (Meiji University)
Siberia and Central Asia: Hirofumi Kato and
Masaki Naganuma (Hokkaido University)
South Asia: Kazuya Shimogama (Ancient Orient
Museum)Southeast Asia: Shinji Yamasaki (Okinawa
Prefectural Museum) and Takehiro Miki (The
University of Tokyo)
East Asia: Kenji Nagai (Tohoku College of Arts
and Designs) and Hiroto Nakata (Aoyama-Gakuin
University)
Comparable Paleolithic databases currently
available include the Stage Three Project database( Van Andel and Davies, 2003), the PACEA
database (D’Errico et al., 2010), and the ROCEEH

database (Haidle et al., 2010). Each database isvaluable in its own right. For example, the Stage
Three Project database focuses on archaeologicalsites of Europe, and as indicated by its name,it concerns sites dating to MIS 3. The PACEA
database also focuses on European sites but limits
its coverage to those that can be dated by theradiocarbon method, that is, 50 ka and younger.
The ROCEEH database is the most extensive,
containing detailed information on archaeologicalsites in Africa and Eurasia beginning at two
million years ago. Partly due to its very large
geographical and chronological scale, this project isplanned to continue for a considerable period.
Similar to the ROCEEH database, the RNMH
NeanderDB covers much larger geographic
areas than the Stage Three Project and PACEA
databases. However, it limits the time range tobetween 200 and 20 ka in order to focus on the
period beginning with the appearance of the first
anatomically modern humans in Africa and ending
with the demise of Neanderthals in Eurasia.
Moreover, NeanderDB contains a detailed list ofarchaeological finds for each assemblage such asbone tools, ornaments, mobile art, and specifictypes of lithic tools (Kondo et al., 2012). Sucha list allows users to examine the invention andmaintenance of patterns of a particular cultural
item across geographical regions so that we canthen test the “learning hypothesis.” Another uniqueaspect of the NeanderDB is that it incorporates
a wealth of information from regions outside of
Africa and Europe such as Siberia, South andEast Asia, and Oceania. Particularly rich are thedata from East Asia, collected from literature in
Chinese, Korean, and Japanese. Clarification of thetime-space context regarding the emergence of theUpper Paleolithic in East Asia, whose indigenous
populations were probably not Neanderthals,
can be compared to the insights obtained fromstudying the Neanderthal record elsewhere.

Palaeolithic Archaeology Group A01

References
D’Errico, F., W. Banks, M. Vanhaeren, V. Laroulandie,
and M. Langlais (2010) Pacea geo-referenced
radiocarbon database. PaleoAnthropology 2011: 1−12.
Haidle, M. H., M. Bolus, A. A. Bruch, C. Hertler, A.
Kandel, M. Märker, N. J. Conard, V. Hochschild,
F. Schrenk, and V. Mosbrugger (2010) The roleof culture in early expansions of humans – A newresearch center. Quaternary International 223-224:
429–430.
Kondo, Y., S. Kadowaki, H. Kato, M. Naganuma, A.
Ono, K. Sano and Y. Nishiaki (2012) Network
computing for archaeology: A case study from the‘Replacement of Neanderthals by Modern Humans’
database project In: CAA2011 - Revive the Past:
Proceedings of the 39th Conference in Computer
Applications and Quantitative Methods in Archaeology,
Beijing, China. 12-16 April 2011, edited by M. Zhou,
I. Romanowska, Z. Wu, P. Xu and P. Vewrhagen, pp.
227-236. Amsterdam: Amsterdam University Press.
Nishiaki, Y. (2012) An archaeological study of thelearning behaviours of Neanderthals and Modern
Humans. RNMH Project Series 1: 32-37.
Van Andel, T. H., and W. Davies (2003) Neanderthals
and Modern Humans in the European Landscapeduring the Last Glaciation: Archaeological Results of theStage 3 Project. Cambridge: McDonald Institute forArchaeological Research.

093

Lithic Refitted Materials and Identification of Technical
Skill Level: Lessons from the Upper Paleolithic Assemblagesof the Shirataki Sites in Hokkaido, Northern Japan
Jun Takakura

Hokkaido University, Japan

Palaeolithic Archaeology Group A01

094

My purpose in this paper is to reassess asignificance of the lithic refitted materials in order
to explore the learning and skill transmission
process in the lithic production. I approach
this issue through the analysis of the enormous
lithic refitted materials obtained from the Upper
Paleolithic assemblages at the Shirataki sites,
Hokkaido, Northern Japan. Importantly the
Shirataki sites are located nearby a huge outcrop of
obsidian that is of good quality, with few interior
inclusions. Therefore, the Upper Paleolithic
inhabitants of the sites procured lithic raw
materials from these local obsidian sources, and alarge amount of lithic production was done at therespective sites. Analyzing and comparing of theselithic refitted materials encourage us to examine
the relationship between the contexts of pastknapping activities occurred at the site and how
skill was expressed materially.
To address such issue, I attempt to assess not
only the technological characteristics which havebeen generally understood as skill signatures in
the previous approaches, but also presence of
blade core as well as the degree of absence of blade
among the refitted materials. The former concerns
whether the blade cores expected to be includedin the refitted materials are recovered from thesite or not. The latter refers a comparison between
the estimated number of produced blades andthe estimated number of missing blades in therespective refitted materials. It is highly probablethat the blade cores and the blades that goodresults were achieved technically were exported
from the Shirataki sites, with an anticipation offurther use as blanks or tools. Conversely, themajority of the products knapped by the novice left
on the spot where they were struck from, which
confirms the hypothesis that a simple training
exercise took place (Bodu et al., 1990). Whether
the blade cores as well as the blades are present
in particular assemblage may provide us with
important information on the skill of knappers andthe context of past knapping activities. Of course,when the accidental flaking seriously occurred,for example, resulting from the inferior inclusions
or the joint surfaces in the obsidian gravels anddebris, and the operational sequences were justterminated by the error, the blade cores should beabandoned at the site. In other cases, the criterion
for abandonment is related to the economic
threshold: work stops when the core only allows
the manufacture of products of insufficient size(e.g., Karlin and Julien, 1994). Without knowing
just how checkered preceding flaking may be, weshould not automatically regard the presence of
blade cores as indicator of the skill level concerning
the prehistoric knappers.
In this paper, I present data documenting theskill learning processes based on the analysis of
the Late Upper Paleolithic assemblages obtained
from three sites in the Shirataki sites: theHattoridai 2 site, the Kamishirataki 8 site, and theKamishirataki 2 site.
First, I attempt to analyze the refitted materials
obtained from the concentrations Sb-23~31 at
the Hattoridai 2 site. A comparison of the refitted
materials shows that a correspondence of the skill
signatures with presence of the blade cores and thethat variability can be seen in the morphologicalfeatures of the raw materials. The evidence
presented here offers that the knappers involved in
the formation of this assemblage had an equivalentskill to produce the blades. They used the various
form of obsidian gravels as raw materials to remove
the blades, which is a contrast to the case in the
Hattoridai 2 site.
In these refitted materials, blade cores are present
except for few specimens. Conversely, the vast
majority of blades supposed to be detached among
these refitted materials were largely exported
from the site. Therefore there is an inconsistency
between them. Interestingly, I can recognize that
remarkable accidental flaking such as hinge fracture
scars and other irregularities usually occurredin the preceding flaking before abandonment
through the observation of these refitted materials,
especially conjoining with the blade cores. Such
accidental flaking might result in the abandonmentof cores at the site. As a result, there are not therefitted materials viewed as reflections of different
levels of technical skill from the concentration
No.90 at the Kamishirataki 8 site. Furthermore, it
is of interest to note that the skill signatures clearlycorrelate with not the absence of blade cores, but
the absence of blades.
Third, I analyze the refitted materials from theconcentration Sb-9 at the Kamishirataki 2 site.
There are four refitted materials relevant to theblade reduction in this assemblage. Each of them,conjoining many of blades and flakes, clearlyshows that how the raw materials were selected,
how preparations and rejuvenations of blade cores
were repeatedly employed, and subsequently how
and what blades were removed. It appears that theblade productions in these refitted materials were
progressed in much the same way. Large chunksof obsidian debris were commonly used as raw
materials in these refitted materials. The diametersof raw materials selected in this assemblage areabout 40 cm. Then, the regular and long blades,from almost 10 cm up to 30 cm in length, were
subsequently aimed to detach among these blade
reductions. To address it, the multistage reduction

Palaeolithic Archaeology Group A01

blades is apparently recognized. Indeed, different
skill levels of the knappers are associated with theselection of raw materials; the skilled knappers
might use angular and sub-angular obsidian
gravels as lithic raw materials. In contrast, theapprentice knappers might use grounded obsidian
gravels for the blade reduction. Consequently,
we can understand that the skill of prehistoricknappers in this concentration was combined withselection of the form of raw materials, regardless ofa consideration concerning an influence from thetechnological needs and behavioral contexts.
It is noteworthy that the distribution of refitted
materials made by the skilled knappers and thenovice knappers is spatially separated in thisassemblage. This is valuable patterning becausesuch distributional pattern perhaps gives a clue toinfer the social relationship between the multiple
knappers with different level of skill. In spite of afew data to claim that the knapping activities inthis assemblage were conducted simultaneously,
a distinction of the spatial patterning enable us
to interpret that the activity zones of the skilled
knappers and the novice knappers are differentiatedbased on well-defined spatial rules, and suchrules therefore might be recognized among bothknappers. This patterning is nevertheless different
from the case of the Paris Basin where reserving
the best places around the hearth for the skilled
knappers and keeping the apprentices outside thedomestic space are seen (Pigeot, 1990).
Second, I focus on the lithic assemblage fromthe concentration Sb-90 at the Kamishirataki
8 site. Careful preparations as well as repeatedrejuvenations on the striking platform and ridge arevirtually recognized in all of the refitted materials.
Although the morphological features of obsidian
gravels vary among these refitted materials, thetechnological characteristics observed in the blade
reduction process and its products are surprisingly
similar. Also, there are a few differences inthe technological evidences in relation to theskill signatures among these refitted materials.
Consequently, it is hard to identify a difference inthe technical skill level of the knappers, in spite of095

sequences including the careful preparation andseveral rejuvenations stages of striking platform as
well as ridge on these blade cores, were executed.
These were always connected with rubbing thepecked platform, presumably with the aid ofabrasives. When the accumulation of hinge fracture
terminations on the face of the cores as well as thestriking platforms occurred, the knappers often
restored to rejuvenating the flaking surfaces andplatforms to eliminate the accumulations of errors
created on these cores. Therefore, the technologicalfeatures, common to all of blade refitted materials
in this assemblage, allow us an interpretation thatthe knapper had the high skill to maintain coresfor producing the formal blades effectively andcope with the errors appropriately.
However, it is of interest to note that manyof blades detached from the blade cores in thisassemblage were abandoned at the concentration
Sb-9, while some of the blade cores were exported
from the site. In these cases, every stage in thecourse of manufacturing processes was performed
with utmost precision, but almost all of the
products were left on the spot. This is in contrast
to the case of the Kamishirataki 8 site, as describedabove. These materials can be interpreted as“academic cores” ( Johansen and Stapert, 2008):
cores worked by an expert knapper in what seems
to be a pedagogic demonstration for the benefit ofa beginner knapper. One conclusion that can be
drawn is that its purpose of the expert knappers
was not to produce good blades for immediate use,
but to instruct.

Palaeolithic Archaeology Group A01

096

Comparison between Individual, Imitative andInstructive Learning

Juko Ando

Faculty of Letters, Keio University, Japan

Methods
Participants:
20 university students (12 females, 8 males); mean
age = 20.8 yrs (SD= .77)
Materials:
Three sets of disentangled puzzles; Puzzle I (Key), II(Horse), and III (Disk) (Figure 1) .
Procedures:
There were three phases; individual, imitative, andinstructive learning.
(i) Individual learning (IndL); To solve the firstpuzzle alone without any assistance by other.
(ii) Imitative learning (ImitL); To solve the second
puzzle in the situation that a learner is permitted
to imitate a model (who learned the target puzzle
in Phase (i)) solving the same puzzle.

Puzzle Ⅰ (Key)

Puzzle Ⅱ (Horse)

Puzzle Ⅲ (Disk)

Figure 1. Materials.

Hunter-Gatherer Studies Group A02

One of the most important research questions
on differences of learning abilities between the
Neanderthals and the modern humans (MH) is
to clarify differences between social learning andindividual learning psychologically. It is evident
that the Neanderthals transmitted their skills
to make stone axes by means of social learning,
especially imitation. The MH could make creative
individual learning as well as imitation. Individual
learning with creativity of the MH is obviously
different from individual learning universally
conducted by non-social animals. Additionally the
MH can make instructive learning as a special styleof learning specific to Homo sapiens (Strauss, 2005;
Csibra and Gergely, 2006; Ando, 2009, 2012).
Although imitative learning is a kind of social
learning in a sense that learning occurs in a social
situation in which models to be imitated exist,
it is also a kind of individual learning because it
depends on subjective understanding of how torepresent models’ skills to be learned and how toregulate learners’ own learning processes. On theother hand, instructive learning is evolutionarily
idiosyncratic because there exist a person who
costly and altruistically control other person’s
learning of learners.
In the current preliminary study, therefore, threedifferent styles of learning - individual, imitativeand instructive learning - were compared to clarify
nature of learning in the MH; how efficient they
are, how motivations to imitate/ to be imitated andto teach/ to be taught are regulated?

097

Figure 2. Learning efficiencies of each phase.

(iii) Instructive learning (InstL); To solve the third
puzzle in the situation that an instructor (wholearned the target puzzle in Phase (i)) is permitted
to teach and a learner is permitted to ask an
instructor to teach how to solve a puzzle.
Three participants built a team and each participant
in a team solved a different puzzle in each phase.
Trial was identified as success when participant could
disentangle two pieces of puzzle and entangle themagain. The number of successful trials within 10
minutes was an index of learning efficiency.
After the Phase (i), feelings of easiness, task
involvement and challenge were asked. After thePhase (ii), feelings of attention to partner andcompetition as a model and learner (imitator) wereasked. After Phase (iii), teaching motivation (as an
instructor) and feeling of being taught directly andindirectly (as a learner) were asked.

Hunter-Gatherer Studies Group A02

098

Results
Frequencies of successes on each phase forthree puzzle tasks is shown in Figure 2. Althoughno significant differences were found for thesefrequencies of each material, individual learningtended to be the most efficient among three learning
conditions.
Person who feel easy on a task tended to pay more
attention to imitators (r=.53) and person who feel
more competitive feeling while imitating tended tofeel more competitive while being imitated (r=.76)
and less being directly taught (r=-.72).

Discussion
Imitative and instructive learning conditions looked
more time consuming than individual learning tosolve these tasks and, for competitive person, were
not motivating to be taught. Task difficulty andmotivation to teach/to taught might be interact witheach other for efficiency.

Elements of Social Learning Among Hunter-Gatherer
and Farmer Children in the Central African Republic

Adam Howell Boyette
Washington State University, USA

Life history theory predicts that the pace andtiming of developmental changes are evolved
features of an organism’s life cycle. Humans arereliant on social learning for survival and it has
been argued that human childhood evolved, atleast in part, because of the opportunity it provided
for social learning for the ancestors of Homo sapiensin our environment of evolutionary adaptation
roughly 200,000 years ago. Thus, it is hypothesizedthat the capacity for cumulative culture, which
has allowed our success as a species, has coevolved
with our lifespan. While much early social learning
consists of cultural information–shared ideas,
beliefs, values, knowledge, and skills held in theminds of other people–culture also patterns humanbehavior and in part determines who children learn
from and in what settings learning takes place.
It follows that, if childhood is an adaptation tolearn skills necessary for survival and reproduction
within an information-dense social environment(i.e. a culture) then children should:
1) seek out and attend to social information andshould be the targets of teaching and social
feedback;
2) spend less time on learning behaviors withage as skills and knowledge are internalized;
3) form social relationships with those who aregoing to be most useful to learn from–those
who children identify with or are similar but
more knowledgeable than themselves.
In this study, these predictions are tested using
data from children in two small-scale societies: the
Aka hunter-gatherers and Ngandu farmers of theCentral African Republic. I examine social learning
processes and the social contexts of learning.

Hunter-Gatherer Studies Group A02

Figure 1. Fractional polynomial smoothing curves of hourly rates of social learning processes by age. Greenindicates Aka, orange indicates Ngandu. Note the difference in scale of the y-axis on each graph. Jitter was added to thescatter plot with a noice factor of 5.

099

Figure 2. Lowess besi-fit lines of work and play rates
during Aka and Ngandu childhood. In general, girls
spend more time working than boys. However, it is clearthat Ngandu children take the initiative during middle
childhood, and work independently of adults. Aka, on theother hand, begin to work alongside adults at this time.

Figure 3. Rates of proximity to same-sex only versus
mixed sex nearest neighbors (social companions).
Lines are lowess best-fit lines.

Hunter-Gatherer Studies Group A02

100

Figure 4. Rates of proximity to social companions ofdifferent classes across ages in both cultures. Orange
indicates only children of the same age category (4-6,
7-12, or 13-16 years old) are within 5 meters of the focal
child. Teal indicates at least one adult is within 5 metersof the focal child. Red indicates that at least one childolder or younger than the focal child is within 5 meters.

Daily Physical Activity and Time-Space Allocationof Pygmy Hunter-Gatherers’ Children in Southeast
Cameroon
Izumi Hagino and Taro Yamauchi

Graduate School of Health Sciences, Hokkaido University, Japan

Introduction
This study aimed to evaluate the physical activity
and daily time-place use of hunter-gatherers
children, and to examine the relationship of these
indices in one of the most traditional populationsin the world.

Results and discussion
The physical activities of participants wereconsidered to be very high. The average of PAL
and steps were above 2.0 and 20,000 for both
sexes (Table 1). Furthermore, the daily PAL wereclassified into “vigorous” in 73% (46/63 persondays) of all observation, and 95.2% (60/63 persondays) of participants walked more than the

Hunter-Gatherer Studies Group A02

Materials and methods
A complete survey was conducted on 44 Baka
children (28 boys and 16 girls, 2-16 years old)
living in the same village (total 86 inhabitants)
in East region of Cameroon during short dry
season. All participants wore a small GPS unit
for conclusive 3 days, and the GPS log data werecollected (44 children, 131 person-days). Fromthe log data, total travel distance (TD: km/day),
activity radius (AR: m/day) and the daily trip
times (hours/day) for each activity points (e.g.
Own village, Forest and River...) were calculated.
Elder 21 children (13 boys and 8 girls) wore an
accelerometer pedometerwith GPS unit for thesame periods, the acceleration and the steps data
were gained, and the physical activity level (PAL)
was calculated.

international recommendations.
As the age increases, TD was generally increased
and AR was expanded (for both indices, r = 0.70
and P< 0.0001), however, a small dispersions were
found in elder boys for both indices. There was astrong positive correlations between steps and PAL(r = 0.82, P< 0.0001), and moreover, TD and AR
significantly increased as the steps increased (TD:r = 0.82, P< 0.0001 and AR: r = 0.64, P< 0.0001).
These results suggested that the physical activity ofhunter-gatherer's children is related to walking.
There was a sex-age differenceamong daily timeplace using (Table 2). In boys, time spent for“forest (1.07 vs. 3.03 hours/day; younger vs. elder)”and “other village (0.05 vs. 1.85 hours/day)” wereprolonged, and the time stayed in “own village(21.6 vs. 16.9 hours/day” was markedly decreased.
By contrast, there was no notable change for timeplace using in girls.
There were significant positive correlation
between the time spent for “forest ” and fourphysical activity indices (TD, AR, steps and PAL).
However, there were negative correlations between
time spent in “village” and these four indices. It isconsidered that going out from the village might
increase children’s physical activities.

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Table 1. Physical activities of participants.

Table 2. Daily time-place allocation (hours/day) by sex-age distribution.

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102

Figure 1. The increment of total travel distance (km/day) by age.

Figure 2. The correlation between steps (steps/day)and time spend in own village (min/day).

Technical Transmission of Hunting Tool Manufacture:
A Case of Spear Hunting Among Modern HunterGatherers in Southeast Cameroon
Koji Hayashi1, Ryota Ishii2, Yuuki Nakamura3, Hideaki Terashima1, andYoshihiro Nishiaki2
1Kobe Gakuin University, Japan

2The University Museum, The University of Tokyo, Japan
3Meiji University, Japan

more than one spear. Six of 11 persons responded
that they had made more than five spears. Of these,
one person had made around ten spears, while theelder had made “countless”.
Regarding the teaching of spear manufacture, 11 of
18 persons responded that they were taught by theirfather and 2 claimed that they were taught by their
grandfather. It is noteworthy that all responders said
they had “learnt by observing” the manufacturing
process rather than from any “direct teaching”. On
the other hand, 8 of 14 persons without inexperienceof spear manufacture answered that they obtained
a spear by gift (donation) from their kinship group.
Thus, it can be observed that kinship serves as a
background to both the manufacture and donationof spears among the Baka. However, regarding actual
spear manufacture, in spite of kinship relations, elders(experienced persons) do not directly conduct any
teaching. It appears that spear manufacture is always
an independent activity.
The first spears tend to be made during adolescence.
At one time, it was considered by Baka men that
there was a symbolic aspect to hunting with spears
during premarital adolescence. On the other hand,when we pursued this with some Baka adolescents,
none of them were experienced in spear manufacture
and only one of them actually owned a spear.
Therefore, we inferred that spears were mainly
important as hunting tools. However, there has
been a spread of hatchets, steel wires and guns as
modern hunting tools. At the same time, in recent
years the frequency of hunting expeditions have been
reduced by deforestation and the establishment of a

Hunter-Gatherer Studies Group A02

In this study, we present fundamental dataassociated with spear hunting among the Baka, who
are modern hunter-gatherers in southeast Cameroon.
Moreover, the spear as a hunting tool was used byNeanderthals. We decided to examine “learning
hypothesis” through looking at functionality, cultural
aspects and the manufacturing process. We focusedon the distribution of manufacturing techniquesamong the Baka and among generations in addition
to the measurement of spears that are in use. In
particular, we noted the nature of educational
discourse and activity relating to these spears andtheir deployment.
Between August 1 and 9 in 2012, we visited fivesettlements and one half-settlement of the Baka
near Lomie in southeast Cameroon. We measured
34 spears and interviewed 34 persons. As a result, wewere able to verify the diversity of spears (size, shape,quality of material, etc.; Table, Figures 1 and 2). No
obvious relationship between the form of a spear andthe game being hunted was found except in relation
to large mammals such as elephant.
Although detailed information about the Baka’s
actual hunting activities was limited, we were able toobtain useful data relating to the age of interviewees
and the manufacture of spears. Among theinterviewees 18 answered that they had experienceof manufacturing spears. Of these, six persons were
producing spears in their “adolescent (wanjo: teens,premarital period)”, four persons were in their 20s,
five persons were in their 30s and one person was inhis 60s (*four persons did not provide an answer).
Eleven of 12 persons responded that they had made103

national park near the Baka’s immediate forest area.
Consequently, a decline in the hunting activities
which use a spear is likely to be matched by adecline in the technical tradition surrounding the
manufacture of spears.
In this study, we also describe and analyse theresults of some interviews with elders regarding thepractice of spear manufacture (Figure 3).
Table: Spearhead types and their frequency at the study villages.

Figure 1. Classification of spearhead types among the Baka.

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Figure 2. Type 2 spearhead.

Figure 3. Spear manufacturing among the Baka in
Cameroon.

“Finding the Spirit to Create” Innovation and Social
Learning Among Aka Adolescents of the Central
African Rainforest
Bonnie Hewlett

Washington State University, USA

risk-taking and exploratory behaviors and; social
valuation of individual creativity (Diamond, 1999:
246).
This research examines the nature, indigenous
understandings, and transmission of innovation
in a hunter-gatherer group. Since little is known
about this topic in foragers, much of the research
is primarily inductive and descriptive. Data werecollected from in-depth interviews and systematic
ranking and sorting techniques with 20 Aka
adolescents of Central African Republic and 5 Aka
adult individuals identified as being “innovators”.
Data included exploration of indigenous concepts
and terms of innovation, characteristic featuresassociated with innovations, and free listing ofindividuals identified as “innovators”. Further
research incorporated free listing of character
qualities of the “innovators”, in-depth interviews
with adolescents seeking out new knowledge (e.g.
innovative behaviors or inventions) and in-depth
interviews with individuals identified as being
innovators. The study tentatively predicted thefollowing hypothesis based upon developmental
and evolutionary theories:
1. Older adolescents will be more innovative
than children and older adults because ofdevelopmental increases in cognitive capacity
and the potential reproductive value ofinnovative behavior.
2. Older adolescent males will be more likely toseek out inventions as they are more likely toengage in risk-taking and exploratory behaviors
and more likely to travel greater distances than

Hunter-Gatherer Studies Group A02

Innovation and social learning are often
described as responsible for playing significant rolesin the evolution of cultural systems with arguments
chiefly structured around the transmission anddiffusion of novel information, behaviors and or
artifacts, the receptivity of a society to inventions
and the role of innovation as an adaptive response
to highly variable climatic or social conditions,providing solutions to new problems and reducing
uncertainty and risk (for definitions of innovationand invention see O’Brien and Shennan, 2010:vi-18; Henrich in O’Brien and Shennan 2010:
99; Kameda and Nakanishi, 2002; Barton et al.,2011). However, not all innovative behavior is in
response to environmental adversity and instability,
nor developed to meet specific needs (Henrich
in O’Brien and Shennan 2010; Diamond, 1999:
246). Inventive behavior is not unique to humans,
and as with social learning, often occurs in avariety of ways and ecologies (Nishida 1987;
Galef, 1990; Tomasello et al., 1993, in Boesch,1995; Boesch and Tomesello, 2003; Kameda andNakanishi, 2002: Lalande and Reader in O’Brienand Shennan, 2010: 37-52). This study is framedaround economic, organizational, and ideologicalfactors influencing the creation, adoption andtransmission of a new behavior, artifact ortechnology, including; societal organization;
proximity to others; social value and prestige
accrued to the innovator; “ownership rights,” tothe technology/behavior (i.e. protection of rewards
for the inventor); indigenous understandings ofthe advantages of the new behavior or technology;

105

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106

females (McDonald and Hewlett, 1999; Hewlett etal., 1986; see also Reader and Laland, 2001).
3. Innovations would be spread by horizontaltransmission, a characteristic form of transmissionin adolescence (peer, intergenerational) (Hewlett
et al., 2002).
4. Older adolescents would pay attention to
prestigious (“successful”) peers from whom tolearn (Henrich in O’Brien and Shennan, 2010:99-120).
Contrar y to expectations, incidences ofinnovative behaviors occurred more frequently in
middle-aged adults than adolescents, with males
more often being listed as innovators than females.
Adolescents of both sexes were more likely toseek out, and pay for, new behaviors, inventions
and technologies than were adults or children.
Both male and female adolescents sought out new
inventions from adults rather than peers, but maleadolescents did not seek out innovations fromfemale adults, while female adolescents sought out
both adult males and females to learn from. Bothmales and females frequently listed being seen as
more attractive to the opposite sex as the reason
for acquiring new behaviors and or technologies.
More data from small-scale populations arenecessary for formulating precise understandingsof innovation, social learning and culturaltransmission in contexts that characterized most of
human history. The consequences of environmental
and biological changes interacting with culturally
driven behavioral changes, innovation, decision
making and social learning may well have beennecessary for the survival of the ancestors ofmodern humans, and help to explain the extinction(or “absorption”) of Neanderthals during the Late
Pleistocene (Barton et al., 2011: 725; see also
Henrich in O’Brien and Shennan 2010).
References
Barton, C.M. et al. 2011. Modeling human ecodynamics
and biocultural interactions in the Late Pleistocene ofWestern Eurasia. Human Ecology 39: 705-725.
Boesch, C. 1995. Innovation in wild chimpanzees.
International Journal of Primatology 16(1): 1–16.

Diamond, J. 1999. Guns, Germs and Steel. New York, NY:
W.W. Norton and Co.
Galef, B. Jr. 1992. The question of animal culture. Human
Nature 3: 157–78.
Henrich, J. 2010. The evolution of innovation enhancing
institutions. In Innovations in Cultural Systems
Contributions from Evolutionary Anthropology, edited by
O’Brien, M. and Shennan, S.J., pp. 99–120. Cambridge,
Mass: MIT Press.
Hewlett B.S., van de Koppel, J.M.H., and van deKoppel, M. 1986. Causes of death among Aka
pygmies of the Central African Republic. In African
Pygmies, edited by Cavalli Sforza, LL., pp. 15-32.
New York: Academic Press.
Hewlett, B.S., Silvertri, A. and Gugliemino, C.R. 2002.
Semes and genes in Africa. Current Anthropology 43:
313-321.
Kameda, T., and Nakanishi, D. 2002. Cost–benefit
analysis of social/cultural learning in a nonstationary
Uncertain Environment. Evolution and Human
Behavior 23:373–393.
Lalande, K. and Reader, S. 2010. Comparative
perspectives on human innovation. In Innovations
in Cultural Systems Contributions from Evolutionary
Anthropology, edited by O’Brien, M. and Shennan,
S.J., pp. 37-52. Cambridge, Mass: MIT Press.
Nishida, T. 1987. L ocal traditions and culturaltransmission. In Primate Societies, edited by Smuts, S.
S., Cheney, D. L., Seyfarth, R. M., Wrangham, R. W.,
and Strusaker, T. T., pp. 462-74. Chicago: Universityof Chicago Press.
MacDonald, D.H. and Hewlett, B.S. 1999. Reproductive
interests and forager mobility. Current Anthropology
40: 501-523.
Reader, S. and Laland, K. 2001. Primate innovation: Sex,
age and social rank differences. International Journalof Primatology 22(5): 787–805.
Tomasello, M., Kruger, A., and Ratner, H. 1993. Culturallearning. Behavioral and Brain Sciences 16: 450–88.

Table 1. Composite salience for Aka attributions ofinnovative people; Males and Females Combined.

Figure 1. Basket old vs 'new'.
Table 2. Composite salience for Aka attributions ofinnovative people; Males Only.

Figure 2. Women's basket.
Table 3. Composite salience for Aka attributions ofinnovative people; Females Only.

Figure 3. Men's basket.

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Figure 4. Drum.

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Hunting Play Among San Children

Kaoru Imamura

Faculty of Economics, Nagoya Gakuin University, Japan

Creation and sharing of images
This study focuses on children’s behavior increating and sharing a new image through playing.
I have carried out a comparative study of children’s
play between Japanese children in kindergarten
and children of the San, who are hunter-gatherers
living in Botswana.
Childhood and children’s play are social andcultural phenomena, as many researchers havepointed out since Phillipe Ariés. Nevertheless,some common points can be seen in children’s play
(Kawabe, 2005):
1. Children will continue to play with concentration
for a certain period.
2. Each child has a certain image about what they are
doing when playing.
3. They voluntarily approach the environment tomake play equipment and the space needed forplaying.
4. They share their play image with others throughmaterials, space and language.

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Observation of children in kindergarten
I have been conducting a comparative study on
children’s play activities at the H Kindergarten in
Seto, Japan, since April 2011.
The results of observations at the Japanesekindergarten are briefly summarized as follows:(1) Playground play
Various types of play activities were observed in
the playground, such as: walking on stilts, playing
soccer, rolling tires, playing tag (cops and robbers),
dressing up in costumes, playing with provided

toys (horizontal bars, climbing ladders, horse-type
toys, jungle gym), playing in the Wendy house or
in dens of large tires, playing with flowers, making“mud balls”, playing in the sandpit, observing
insects in a box.
(2) Sandpit play
In Japan, “sandpits” are so popular for children’s
play areas that every kindergarten, nursery schooland park for children has a sandpit. By playing ina sandpit, children develop their imagination andcreativity. In sandpits, children make “sand houses”
and “mountains,” and also dig holes for “rivers
and lakes,” so that they can make a “town.” Not
only do they play at building things, they also playhouse by imagining the sand as “food.”(3) Sharing images
The children who are playing together in thesandpit share the images of the “towns” and“landscapes.” It is important to clarify how they
can share these images.
(4) Transmission of skills from children to children
In that kindergarten, children make “mud balls”
from sand, soil and water. The teachers do notteach how to make these, the skills are transmitted
from older to younger children.
A field survey of children’s play and learning
among the San in Botswana
I conducted a field survey of children’s playamong the San in Botswana from 11 February to 7
March, 2012. The research points are as follows:
1. Variations of children’s play and the sex-age
composition of their games.

Around ten years ago, children aged four to 14
played hunting games. They had little bow andarrows and they actually hunted little birds andlizards. Everything was small and not serious. So it
was a kind of play.
Now they mimic the whole process of hunting
on horseback. First they make horses and spears
from tree branches and leaves. Next they take theroles of “hunter on horseback” and “game animals.”
Then they perform each role. The “hunters” (usuallyolder children) chase down the “game” (usually
younger children) and they grapple and fight for a
few minutes. Finally the “hunter” kills the “game”
with the “spear.”
They also mimic butchering the game animal.
They reproduce the actions of cutting the “skin”,
then skinning and jointing the animal.
Recently real hunting activities have declined
and the chances for children to observe hunting
have also been reduced. They reproduce hunting
scenes from hearing adults talk about past hunts
and observing the butchering of cattle. They devise
new types of play and share them. Finally theycreate a new imaginary world.
Reference
Kawabe, T. 2005 Nurturing Centered by Play,
Houbunshorin, Tokyo (in Japanese).

Hunter-Gatherer Studies Group A02

2. Transmission of skills for hunting and gatheringfrom older to younger children.
3. Comparing the San children’s “play imitating
adults or others’ behavior” with Japanese children’s.
In this research I focused on mimicking behavior.
In general imitation has a model and imitation
is a behavior in which a person copies the model.
But often children’s imitation play does not needa model. Their imagination can go beyond themodel. Sometimes they share their imaginations
and create a new kind of play. I divide imitation
into two types.
(1) Copying the modelThis type of imitation is found in transmissionof techniques. When children learn how to use aknife and how to make a bow and arrow, a spear, ora snare, they observe adults who are making thesethings and they copy the adults’ actions exactly. But
it is not necessary for the children and adults (themodels) to be together on the spot. Sometimes
children will make a snare alone after they haveobserved adults’ behavior and remember it.
Adults do not directly teach children how to
use a knife. It is the older boys who transmit knife
technique to the younger ones. Neither do theolder boys specifically teach the technique but
they help the younger boys when they reach an
impasse in making some hunting tools or toys. The
way of “helping” is simply an older boy alternates
making them himself and continuing to makewhat a younger boy has begun until it is complete.
This “alternating maker” is often seen even among
adults. They say “alternating making” is “enabling
the owner (who first began to make it) to makesomething.”(2) Sharing imagination and creating new play
“Gokko asobi” (in Japanese) is typical of thisimitation play. Gokko asobi or “pretend play”
involves imitating someone, usually a mother, afather and a baby, while doing role play with otherchildren. The model might be their own family, but
often children act out a “typical” or “ideal” family.
It is a stereotype that goes beyond the real family.
This year, I found a new type of play among San
children. It is “hunting play.”109

Play Activities and Cultural Transmission Among
Hunter-Gatherer Children: The Case of the Baka in
Cameroon
Nobutaka Kamei

Aichi Prefectural University, Japan

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110

Purpose of the research
In order to clarify the causes of the replacement of
Neanderthals by Homo sapiens in aspects of culturaltransmission and culture change, it is necessary to
examine the differences between learning behaviors
and abilities of Neanderthals and those of Homosapiens.
It has been pointed that hunting-gatheringsocieties have neither educational institutions
nor educational behaviors among their children.
Hunter-gatherer children have their own culture,
which is connected to the transmission of huntinggathering culture. These were the results of myanthropological 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 ofschooling and the introduction of restrictions
on hunting, how have children’s culture and lifechanged? Are the characteristics of culturaltransmission in foraging societies still maintained?
This research aims to collect ethnographic data onchildren’s daily activities including play, hunting,gathering, fishing, housework and so on. It also
aims to describe the system of cultural transmission
through this age group as one of the characteristics
of foraging societies.
Methods[Period] March 2012 (5 days); August-September
2012 (14 days)[Field site] Settlements of Baka hunter-gatherers in

the East Region, Republic of Cameroon (Figure 1)[Methods] Participant observation and interviews
with Baka children. Experimental methods onlearning and teaching 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 schoolingis already shared among the majority of parents(Table 1).
(2) Play and subsistence activities (hunting,gathering and fishing) conducted by boys and girls
observed out of school have not changed.
(3) Social and cultural changes among adults partly
affect children’s activities, but do not affect their wayof learning.
Discussion
The results show that in the past fifteen years,
children’s culture and life have not changed.
Observation and imitation are still the essential
ways of learning among children. As before, it is rare
to observe positive teaching activities among them.
The characteristics of foraging societies “without
education” are still maintained.
These comparative data show us both robust andvulnerable aspects of the way of life and behaviors
of hunter-gatherer children. Together with theseobservations, a “learning model of hunter-gatherer
children” will be presented in order to discuss thenature of “educational activities” by human beings.
It is hoped that this will contribute to clarifying thecauses of culture changes and innovations that led
to the replacement of Neanderthals by Homo sapiens.
Kamei, N. 2010. Little “hunters” in the forest:
Ethnography of hunter-gatherer children. Kyoto:
Kyoto University Press.

Figure 1. Field site.

Table 1. Number of Baka children of school age (6-12 y.o., March 2012).

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111

Cognitive Flexibility and Making Objects in Baka
ChildrenTadashi Koyama

Faculty of Humanities and Sciences, Kobe Gakuin University, Japan

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112

Relying on Piaget’s theory, Pellegrini (2009)
pointed that children’s cognitive development
could been rooting in their sensor imotor
interactions with objects. Through repeatedinteractions with objects, children constructrepresentations for those objects (Pellegrini, 2009).
These representations would come to express intheir making objects. Some aspects of children’s
cognitive development and their ongoing interests
are reflected in their making objects. And doing inthese activities may develop their creativity. Child’s
behavior with objects has been studied from theview point of exploration, play, construction, andtool use (Bock, 2005). Children would often beabsorbed in making objects for their play, suchas making a basket out of leaves. I think these
activities affect later their creativity and cognitive
flexibility.
I n re c e n t re s e a rc h e s i n d e ve l o pm e n t a l
psychology, cognitive flexibility is defined that achild could switch his strategy to solute the tasks.
For example the DCCS (Dimensional Change
Card Sorting; Zelazo, 2006) task assesses thecapacity to flexibly shift between two sorting
rules based on color and shape. Kloo, Perner,
Aichhorn, & Schmidhuber (2010) suggested that
we had to distinguish two different kinds, response
flexibility and representational flexibility. Making
objects also requires children’s cognitive flexibility,
especially representational flexibility. I want todemonstrate the creativity of the children who are
nurtured in hunter-gathering societies. So in thisstudy I observed Baka children’s making objects
in their daily life and speculated on their cognitive

flexibility through their making objects.
1. Method
I conducted the observational research on
children’s making objects in East Cameroun
between 13th and 18th of August in 2012, I
visited five Baka Pygmies villages and conducted
observation on Baka children’s making objects andplaying with objects in their site. I observed about
two hours in the morning and two hours in theafternoon every day. The children’s activities with
objects were recorded by a SONY Handy Cam.
2. Results
Making objects for play were very often observedin Baka children. But the scene of making objects
was observed almost in solitary play (Figures 1,
2 and 4). And making objects and constructing
objects were observed in collaborative symbolic
play (Figure 3). This play was observed particularlyin the girls. They used object substitution for many
purposes to simulate the real cooking scene. Somechildren incorporate ‘newcomer’ materials to theirsites into making objects for play (Figures 1 and 2).
Those products then were used in other children
in the same site. And it came to be elaborated bythem. Figure 2 shows that the boy used concave
space on the ground as pretending a river bridged
it with wood. It seemed to represent the real scenethat a car with loading went on the bridge.
Figure 4 shows solitary constructing objects of 4
year old boy. The boy in Figure 4 was concentrated
in attaching some pieces of wood to the wheel.
Children would use those broken parts of entitiesfor their objects making. Figure 5 shows that theboy in the picture making a steering while using
broken parts of a motorcycle.

Figure 1. Making an object with acan.

Figure 2. Making river and its bridgewith wood.

Figure 3. Making objects in girl’scollaborative symbolic play.

Figure 4. Constructing objects with awheel and wood.

Figure 5. Using broken parts of a
motorcycle.

Hunter-Gatherer Studies Group A02

3. Discussion
Until about 3 years old, children engaged insensorimotor interactions with materials aroundthem. Around 4 years old, children would makeobjects. But these observations reported here werevery often and making objects were observed
almost in a child’s solitary situation.
Sex differences were also observed. Girls making
and constructing objects were observed in theircollaborative symbolic play, which was similar tocooking and preparing meals that woman engaged.
Those activities were interpreted playful uses ofobjects as simulations. And object substitution inthat play might increase their cognitive flexibility.
On the other hand, boys preferred making objects
for pretending as cars, trucks and motorcycle. For
that activity new materials tended to incorporate
with leaves, stones, wood, and string. It wasthought that in that construction of object their
cognitive flexibility may be enhanced.
Kamei (2009) reported materials that Baka
children used for their play. And he emphasizedthat ’s richness and creativity of them using
those materials. Kloo, Perner, Aichhorn, &Schmidhuber (2010) demonstrated that‘Flexibility
is a key competency in daily life.’ We can see
Baka Pygmies children’s development cognitive
flexibility in their daily making objects, object
substitutions and using broken parts of entities.
In this research I noticed new materials might
affect their development. Koyama (2012) pointed
that younger Baka children showed their intuition
and ability of representation in responding tosigns of visual reception tasks. The developmentof representation in responding to signs of visual
reception tasks parallels that of making objects.
Intuition and ability of representation might berelated to the development of their object making.
This research impressed on me that children’sdevelopment of cognitive flexibility are based on
their intentionality to real world around them.

Acknowledgments: This research was supported
by Grant-in-Aid for Scientific Research on
Innovative Areas 2010-2014, 1201, Project
leader Takeru Akazawa, PhD., Replacementof Neanderthals by Modern Humans: Testing
evolutionary Models of learning.

113

The Art of Patience as the Emotional Basis for
Creativity and Cultural Learning: Learning andEcological Adaptation Among the Canadian Inuit
Keiichi Omura

Graduate School of Language and Culture, Osaka University, Japan

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114

Any human individual is born into each society
and begins its lifelong learning process with social
learning of each culture. In this sense, social learningof cultures can be said to be the base for any kindsof learning for Homo sapiens.
Above all, as pointed out by Tomasello (1999),‘cultural learning,’ including ‘imitating learning,’‘instructed learning’ and ‘collaborative learning,’
plays the most important role in the learning
process of Homo sapiens. This is because culturallearning is essential to its species-unique modes ofcultural transmission, that is ‘cumulative cultural
evolution,’ the process of which ‘requires not only
creative invention but also, and just as importantly,
faithful social transmission that can work as aratchet to prevent slippage backward - so that the
newly invented artifact or practice preserves its new
and improved form at least somewhat faithfully
until a further modification or improvement comesalong.’ (Tomasello, 1999: 5) If so, how does humanindividual learn each culture and improve what s/he
has learned by cultural learning?
In order to understand the mechanism of‘cumulative cultural evolution,’ I conductedethnographical research on the learning processof Inuit children at Kugaaruk, Nunavut, Canada,
between the 8th and 27th of March in 2011,
between the 12th and 27th of March and between
the 17th of August and the 7th of September in2012. The result of these researches is as follows.

Kugaaruk, Nunavut Territory, Canada

one of extended families in Kugaaruka teenager withhis harpoon

narwhale hunting

uncle and niece

a girl learning the wayto
dissect narwhale

adults and children at homea boy hunting caribou

2) Inuit children are exposed to adults’ playful
teasing in everyday life and learn the art ofpatience as the base for cultural learning andcreative individual learning through theirexperience of being in a dilemma posed by theteasing.
3) It is crucially important to investigate how Inuit
adults deal their children in their everyday life
in order to understand how Inuit children learn
the emotional attitudes essential to flexible
and creative individual learning and culturallearning.
Reference cited
Tomasello, M. (1999) The Cultural Origins of Human
Cognition. Harvard University Press.

Hunter-Gatherer Studies Group A02

1) The flexible and creative tactics, which
are considered to be the essence of Inuit
Qaujimajatuqangit (Inuit Knowledge), arelearned by experience of overcoming difficulties
with persevering and patient efforts in everyday
life and subsistence activities.

115

Communicative Musicality and Learning in CaregiverChild Interactions Among the San of North-Central
Namibia
Akira Takada
Kyoto University, Japan

In this paper, I will discuss the role of musicality
for promoting early socialization and learningin the course of caregiver-child interactions
among a group of the San (!Xun), a post-huntergatherer people living in the north-central partof Namibia. Musicality introduces the structureof “intersubjective time” (Gratier & ApterDanon, 2009) as a means of achieving mutual
understanding in caregiver-child interactions.
This facilitates cooperative coordination of
attention, posture, position, and affect among
participants. This coordination communicateswhat the participant should and should not doand thereby provides children with the ground forsocialization and learning in the given society. This
process prepares a context in which actions that fit
with culturally shared values are put into practicein everyday life. The communicative musicality
perspective (Malloch & Trevarthen, 2009) give apowerful tool to analyze how such process is going
to be organized.

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116

References
Gratier, M. & Apter-Danon, G. (2009) The improvised
musicality of belonging: Repetition and variationin mother-infant vocal interaction. In S. Malloch& C. Trevarthen (Eds.), Communicative musicality:
Exploring the basis of human companionship (pp. 301327). Oxford University Press.
Malloch, S. & Trevarthen, C. (2009) Musicality:
Communicating the vitality and interests of life. In
S. Malloch & C. Trevarthen (Eds.), Communicative
musicality: Exploring the basis of human companionship(pp. 1-11). Oxford University Press.

The Resilience of Pygmy Children– On Their
Drawings about Favorite Things

Eiko Yamagami

Kobe Gakuin University, Japan

Introduction“Resilience” is a key word for understanding how
to cope with extreme stress such as the trauma ofchild abuse or serious experience of war and disaster.
In fact, although some victims suffer from PTSD
and some of them develop a personality disorder
after growing up, some of them survive without
any problems. The last group might be successfully
surviving as they are flexible and capable of learningfrom their difficult condition. The strength of theflexible attitude is “resilience”, or in another word
willowy. This psychological concept “resilience” might
be applied for considering the superiority of Homosapiens, because of their ability to overcome a difficult
environment, including temperature, food and enemy
issues over a long history. I suggest that the resilienceof Homo sapiens, examining the drawings of huntergathers Pigmy Children who might be a pure type of
Homo sapiens.

Procedure
The children were asked to draw their favorite
things on the paper with tools such as pencil, colored
pen and pastel crayon, which they could choose.
Results
1) Drawing process and attitude
They chose paper and tools that were set up in

Discussion
The favorite things drawings of Pigmy children
seem to reveal the resilience through their drawing
process and drawing itself. For example, one of theresilience factors, “Productive activity” (Klohnen,1996) might be shown by their active attitude foran unfamiliar situation and kinetic and energetic
drawing style. The drawing content also indicates

Hunter-Gatherer Studies Group A02

Participants
31 Baka Pigmy Children in Cameroon (25 boys
and 6 girls, age ranged from 5 to 15 years old, thedata collected in 2011 summer)

front of them. Then, according to my instruction, they
started to draw spontaneously without hesitation or
help from others. After making images they pointed
and named their favorite things on the paper. Finally,
they told me that they had enjoyed drawing verymuch.
2) Drawing itself
Drawing style: 1 constructed painting (Figure1), 15 imaginary x-ray paintings (Figure 2), 10
incomplete gestalt paintings including 2 head-feet
representations (Figure 3), 17 multiple viewpoint
paintings, 18 kinetic paintings (Figure 4), 18 energetic
paintings and 2 skillful and artistic paintings (Figure
5).
Drawing content (rate of drawing): vehicle andplane 80.6%, human being 74.1%, food (edible
animals and fruits) 64.5%, plants 61.2%, houseincluding a traditional tent mongle 38.7%, tools(pump, rifle, traditional knife, drum, pan, umbrella,glasses, wooden boat and table) 38.7%, animal 32.2%,
fish 32.2%, soccer ball 29.0%, electrical appliances(T.V., DVD, mobile phone, antenna etc.) 19.3%, sun,moon and stars 19.3%, letters and numbers 16.1%,
cloth 6.4%, and national flag 6.4%.

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“Productive activity”. Namely, “shot with rifle” and“kick a soccer ball” in the drawings are important
activities for their life. “Hand” is needed to attainsomething and “food” is the desire to eat forsustaining life. All these figures might reveal“Productive activity”. Moreover, the content showsother component factors of the resilience. For
example, a high rate of “human being” might mean
friendship with others or staring at themselves andso it might indicate the resilience factor, “Insight andwarmth”. “Vehicle, plane and electrical appliances”,
which belong to a different culture, are far from thereal life for Pigmy children, but they are fond ofsuch unfamiliar things and expect to obtain them infuture. The children can imagine how the new andHunter-Gatherer Studies Group A02

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strange things are desirable. Likewise, they are infavor of both their own and different culture related
things in their drawings. This openness might mean
the resilience factor, “Confident Optimism”. The lastresilience factor by Klohnen, “skilled expressiveness” isshown by the art activity for all of them, especially, by
two skillful and artistic paintings.
Conclusion
Pigmy children can be considered to be resilient
people. Through their “resilience”, i.e., an acceptableand flexible attitude to a transitional and criticalsituation, I speculate that Homo sapiens have been
thriving in the face of adversity over a long history.

Figure 1. Constructed painting. A (9yr. boy).
Car with antenna, house, person, lizard, dog,mouse, trees, rifle and birds

Figure 2. X-ray painting. B (10yr. boy). Car,house, person, ball, pump, hand, fish, cook,
tomato and flower

Figure 3. Incomplete gestalt painting.C (11yr.
girl). House, persons, cook, car, table and tree

Figure 4. Kinetic painting. D (7yr. boy): Manwith rifle, persons, cook, car, cacao, hand

Figure 5. Artistic painting. E (12yr. boy).
Helicopter, motorbike, radio, DVD, ball, boat,
mobile phone and moonEstimation of the Period of Childhood and Child
Growth Characteristics of Pygmy Hunter-Gatherersin Southeast Cameroon
Taro Yamauchi and Izumi Hagino

Graduate School of Health Sciences, Hokkaido University, Japan

Introduction
Pygmy hunter-gatherers, who are known as theoriginal inhabitants of Central African rainforest,
have the smallest stature in the world. Their shortstature had been attracted by many anthropologists
and human biologists, and so their growth pattern
as the mechanism of their physique was thought as
the important theme of human biology which canprovide key knowledge of human evolution. This
study aimed at characterizing the growth patternof pygmy hunter-gatherers by using mathematicalmethod and comparing with world populations.

Results and discussion
The adolescent growth spurt of Baka children
were characterized as strongly suppressed thanpreviously studied children (Figures 1 and 2; solidbold lines).
The period of adolescent growth spurt, which
was indicated as from age at take-off (ATO) topeak height velocity (PHV ) was almost similar
among all groups for both sexes (Tables 1 and 2).
It is considered to be reflected the good nutritional
status of Baka children as well as over 80% ofparticipants had normal ranged BMI. This result
showed that the periods of adolescent and also theperiod of childhood were similar among modern
human.
There was a remarkable characteristic about
growth tempos of Baka children during earlychildhood. Although adult height were greatly
difference among Baka and other groups both
sexes (boys: 11.6 - 20.5 cm, girls: 4.1 - 15.8 cm),the amounts of height gain after the onset ofadolescent growth spurt were quite similar amongsix populations including Baka children (Tables 1
and 2). It may suggest that the height gain during
early childhood is much less, and the growth
tempo of Baka children was slow.

Hunter-Gatherer Studies Group A02

Materials and methods
A cross-sectional anthropometric measurements
including height and weight were conducted on
626 Baka children (349 boys and 277 girls) from62 semi-settlement villages ranged wide area insoutheast Cameroon. The ages of participants
were estimated to the nearest 0.5 years by birthreordering, interviewing and event matching.
The smoothed growth and height velocity curves
were derived from applying Preece-Baines modelI, which is one of the most common mathematicalhuman growth models. The biological parameters(e.g. ATO: the onset of adolescent growth
spurt, PHV: velocity at peak height velocity)were calculated from mathematical parametersof function, and compared with 5 populations(Europe, Africa, Asia, Australia, South America)
from previous studies.
The nutritional status which influences thechild growth, was assessed by classification ofparticipant’s BMI using sex-age cut-offs proposed
by International Obesity Task Force.

119Figure 1. Height velocity curves of boys.

Figure 2. Height velocity curves of girls.

Table 1. Biological parameters of Baka compared to previous studies (boys).

Table 2. Biological parameters of Baka compared to previous studies (girls).

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The Effects of Cross-Boundary Rituals on Cultural
Diversity

Shiro Horiuchi

Shibaura Institute of Technology, Tokyo, Japan

Introduction
In this talk, I examine how cross-boundary
rituals, through which members of different groups
communicate with one another, affect cultural
diversity in modern human groups. In general, massmigration destroys local minority culture, which
may also have been the case in prehistoric times.
Cross-boundary rituals should have counteracted
the negative effect of migration and maintained thediversity of local cultures. To test this hypothesis, I
built an agent-based model (ABM) simulation. Thismodel is a different version of the one described by
Horiuchi and Kubota in an oral presentation.

Results
When L = 0, the value of D produced a decreasing
curve along the value of log Q. When L = 1, the valueof D showed a U-shaped curve along the value of log
Q; the values of D were significantly higher when
agents migrated more often (1 ≥ log Q ≥ -2) thanwhen agents migrated less often (5 ≥ log Q ≥ 4) (post
hoc Kruskal-Wallis test, P < 0.001). Mass migrationsof agents left fewer cultural traits when agents did
not carry out cross-boundary rituals; more cultural

Evolutionary Models Group B01

Methods
The ABM assumes N agents and M groups. The
groups are arranged along circular stepping-stones.
N / M agents belong to each group under the initialcondition. Culture consists of K independent traits,
represented as the vector (ci, 1, ci, 2, …, ci, K) for
agent i. If the agent knows, or does not know, the jthcultural trait ci, j is 1 or 0, respectively. Ci denotes
the total number of cultural traits that agent i knows,
which can be expressed as Ci = ∑jci, j. D denotes thetotal number of remaining cultural traits which are
known by at least one agent. The range is 0 ≤ Ci, D≤ K. At the initial condition, agents of group j know
only the jth cultural trait, so Ci = 1 and D = M. Herecultural trait j is the endemic knowledge of groupj. For simplicity, the model sets parameters as N =200, M = K = 20. The total number of groups equals
the total number of cultural traits, and the set of allgroups is matched against the set of all cultural traits.

The simulation iterates turns composed of two
steps: a within-boundary ritual and a cross-boundary
ritual. In the within-boundary ritual, two agents inthe same group are selected. One copies a randomly
selected cultural trait from the other. In the crossboundary ritual, an agent checks her own group andits nearest 2L groups (0 ≤ L ≤ M/2). Among these,
she selects the group with the largest average valueof Ci. She then visits that group, copies a cultural
trait from an agent in that group, and goes backs to
her own group. In these two processes, agents maylearn or forget a cultural trait, when counterparts do
or do not know the trait. An agent migrates between
groups in Q turns (10-2 ≤ Q ≤ 106). When Q issmaller than 1, 1 / Q agents migrate between groups
for each turn.
I used the parameter D as the index of cultural
diversity. The purpose of this simulation was toinvestigate the average value of D for 30 trials undereach condition of (L, Q) after enough time passed(preliminary simulations indicated that 300,000 turns
were sufficient).

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traits remained due to mass migrations when agentsdid carry out cross-boundary rituals (Figure 1).
When log Q = 2, the value of D increased along
the value of the L curve. When the value of log Q =0, the value of D showed an inverse U-shaped curvealong the value of L; when L = 10, the values of D
were significantly smaller than when 1 ≤ L ≤ 9 (post
hoc Kruskal-Wallis test, P < 0.001). The greatest
cross-boundary rituals did not always maintain highcultural diversity when agents frequently migrated
between groups (Figure 2).

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122

Conclusion
If agents perform within-boundary rituals only, andmigrate between groups, each cultural trait should
disappear from the world by a probability of 19/20
once enough time has passed. On average, only one
cultural trait should remain (D = 1). More cultural
traits will remain in the world if cross-boundary
rituals are also performed. It is presumed that
Neanderthals maintained a monoculture over thewhole European continent, whereas modern humans
propagated cultural diversity. Different local cultures
were sometimes united and evolved into a moreelaborate culture. In both Homo species, individuals
migrated between groups. But only among modern
humans were cross-boundary rituals enacted. Crossboundary rituals could have affected the culturalaccumulation of modern human groups and resulted
in their replacing Neanderthals.

Figure 1. Without cross-boundary ritual (L = 0), few
cultural traits remain (small D) when agents frequently
migrate between groups (small log Q). With crossboundary ritual (L = 1), more cultural traits remain (large D)
if agents frequently migrate between groups (small log Q).

Figure 2. When agents frequently migrate between
groups (log Q = 2), more cultural traits remain (large D)
when many groups engage in cross-boundary ritual (large
L). When agents frequently migrate between groups (logQ = 0), fewer cultural traits remain (small D) if all groups
operate cross-boundary ritual (L = 10, the completegraph).

Effect of a Heterogeneous Environment on Range
Expansion of Individual and Social Learners

Kohkichi Kawasaki

Faculty of Culture and Information Science, Doshisha University, Japan

In the previous paper (Wakano et al., 2011),
we investigated the spatial population dynamics
of individual and social learners by a reactiondiffusion model. In the model, it is assumed thatsocial learners may fail to find a conspecific fromwhich to copy the adaptive behavior when the
population density is low. By analyzing the model,
we have following results. When individual andsocial learners are expanding into empty space,traveling waves are formed. Although the space isenvironmentally homogeneous, individual learners
can survive near the front of the traveling waves.
In this paper, we consider the effects of spatial
heterogeneity on range expansion of individual
and social learners by the same type of theprevious model except the space is environmentally
heterogeneous. The space consists of different typesof environments, each other (Figure 1). In theheterogeneous environment, correct social learners
become wrong learners by moving other type ofenvironments. Individual learners quickly learn andadapt their behavior to new type of environment.
Therefore, for the individual learners it is likelythat the environment is uniform.
where Xi is population density of social learners
which behave correctly at type-i environment, Y is
population density of social learners which cannot
correctly behave at everywhere and Z is population
density of individual learners. r is the growth rate,
A is the attenuation rate, m is the competition
coefficient, s is the mortality cost of wrong
behavior and c is the exogenous cost of individual
learning, and D is the diffusion coefficient.
After analyzing the model, we can get results thatthe population of individual learner can be easily
maintained in heterogeneous environment thanuniform environment at the front of expanding range.
Reference
Wakano, J.Y., Kawasaki, K., Shigesada, N. and Aoki, K.,
2011. Coexistence of individual and social learners
during range expansion. Theor. Popul. Biol. 80: 132–140.

Mathematical model is as follows:
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123

Figure1. Heterogeneous environments.

Figure 2. Numerical result for the invasion of social
and individual learners into empty space. r=m=D=1,
c=0.16, s=0.5 and L=20. Solid line: social learner
behaving correctly. Dotted line: social learner behaving
wrongly. Break line: individual learner.

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124

Dispersals of Modern Humans: Inferences from
Genome Diversity Data

Ryosuke KimuraTransdisciplinary Research Organization for Subtropical and Island Studies (TRO-SIS),
University of the Ryukyus, Japan

Introduction
We face exciting times in population genomics.
Because of advances in DNA technologies,
in particular, DNA microarrays and nextgeneration DNA sequencers, we have revealed
genetic structures of human populations at an
unprecedented resolution. In addition, various
statistical techniques have been developed so
far to infer demographic history of populations
from genomic data. However, it is still difficult toreconstruct a complicated demographic history of alarge number of populations. In this study, I focusedon 1) phylogenetic analysis, 2) principal component
analysis (PCA), and 3) model-based clustering
analysis, and aimed to learn how to interpret theresults of these genetic analyses using computer
simulations. Such empirical understandings ofstatistical methods must facilitate to set a complex
demographic model to be tested.

Results and Discussion
Phylogenetic analysis, PCA, and clustering analysis
each shows some specific patterns depending on
the demographic scenarios (Figure 1B-D). In
addition, I demonstrated a possibility that evenmathematical artifacts in these analyses can give acue for discriminating scenarios such as population
split, migration, and admixture. In recent population
genetics, the model-based Bayesian approach is
widely adopted to know the complex demographic
history, in which we need to try a model for
testing its validity. If there are a number of focal
populations and thus a number of possibilities, it
may be difficult to find the appropriate model. Evenin such a case, the statistical methods taken up in
this study, compensating each other, can play crucial
roles in the modeling of possible demographic
scenarios. The empirical understandings of thesestatistical methods will be useful to pare down
possible candidates of the past demographic history.

Evolutionary Models Group B01

Methods
Coalescent simulations assuming multiple models
were performed using a program, msms. We set anin-group consisting of nine subpopulations (SP19) and two out-groups (O1, O2) in the following
models (Figure 1A): cascaded splits model (S),migration model (M), old admixture model (OA),
and recent admixture model (RA). We sampled 20
sequences from each out-group or subpopulation,and obtained 220 sequences in total. The numberof replications was more than 10,000 times. One
SNP with the minor allele frequency no less than
10% was randomly collected for each replication,

and we generated 220 haploid genome data that
contain 10,000 independent SNPs. Finally, two
haploids were paired to construct diploid genome
data. Nei’s minimum genetic distance was calculated
between every pair of populations. From the
distance matrix of populations, we constructed NJ
tree and NN network using SplitsTree4, althoughthese phylogenetic analyses can be applied to the
distances between individuals. Individual genotype
data were applied to PCA using Eigensoft 3.0. A
model-based clustering analysis was also performed
using ADMIXTURE.

125

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126

Figure 1. Simulation study. (A) Simulation models. (B-C)
Results of PCA with outgroups (B) and without outgroups(C). (D) Results of clustering analysis.

A Simulation Study on the Replacement of Neanderthals byModern Humans in Europe: Implications of Climate Change,
Cultural Diversification, and the Shape of the Continent
Yutaka Kobayashi

Department of Biological Sciences, The University of Tokyo, Japan

species, which I call for simplicity Neanderthals
a n d m o d e r n h u m a n s . E ac h l a n d a re a c a n
accommodate up to N human bands. Each band
may bear up to n distinct techniques useful inforests and likewise up to n techniques useful inplains. In every time step, the following events takeplace in order: Markovian vegetation transitions,reproduction, death of bands, and innovation andloss of techniques. In the reproduction phase, eachband produces a daughter band with probability
B. The mother band stays in the current area.
The daughter band, on the other hand, stays in
the natal area with probability 1-m and disperses
to each of eight adjacent areas (i.e. Moore
neighborhood) with probability m/8. The daughterband successfully settles with probability 1-x/N,
where x is the number of preexisting bands in thedestination area. The daughter band inherits all
the techniques of the parent band. In the deathstage, each band dies with a probability dependent
on the local temperature and the technologicalstate of the band. I assume that the death rate ofa band decreases with the number of techniquesthat are useful in the current environment. In
the stage of innovation and loss of techniques,
each band may lose techniques with a constantrate. I suppose, for simplicity, that Neanderthals
are very conservative, so that they never forgettechniques. Likewise, modern humans never forgettechniques that are currently in use. However, they
may forget techniques that are currently not inuse. For simplicity, I suppose that Neanderthals
are not able to innovate new techniques. Modern
humans innovate useful techniques (e.g. forest-

Evolutionary Models Group B01

It is said that Homo sapiens appeared in Africa
roughly 200k years ago and replaced Neanderthals,
who had been occupying Europe, roughly 40-30k
years ago. There are a variety of hypotheses as tothe cause of the replacement, and some of thememphasize the roles of climate change, competitive
exclusion, and inherent difference in learning
abilities between the two species (the “learning
hypothesis”). As is well known, the rate of cultural
evolution and cultural diversity apparently
increased in the Upper Paleolithic in which thereplacement supposedly occurred. In addition, it
is said that Neanderthals survived longer in the
Iberian Peninsula than in other areas of Europe.
In this study, I construct a spatially explicit model
to simulate the replacement of Neanderthals
by modern humans based on multiple causes,
including climate change and difference in learning
abilities.
I consider a virtual lattice space consisting of
80x80=6,400 cells. The cells are categorized into
water and land areas, where the latter are further
categorized into forests and plains. The vegetation
state of each land area stochastically changes
with the course of time following a discrete-timeMarkov process. For simplicity the temperature
is assumed to depend on latitude but not on
longitude. I assume that the transition probability
from a forest to a plain decreases with temperature,
while the probability of the reverse transition
increases with temperature. The temperature varies
cyclically with time, imitating highly variableglacial climate.
I consider population dynamics of two virtual127Evolutionary Models Group B01

128adapted techniques when in forests) at a constantrate. To simulate the replacement of Neanderthals
by modern humans in Europe, the distribution ofland areas was generated from a real map of theworld. Neanderthals were initially distributed over
mid-latitude areas with density 0.1, and modern
humans over low-latitude areas with the samedensity.
Simulations have shown that the replacementof Neanderthals by modern humans typicallyproceeds in three distinct phases (Figures 1 and
2). In the first phase, modern humans cannot
travel north beyond a certain latitude because
Neanderthals basically fare better in such cold
areas. Since the distribution of modern humans
is mostly confined within permanent forest areas,
their population does not fluctuate much despite
variable climate. Neanderthals, on the other hand,suffer from rapid vegetation shifts associatedwith climate change, and their population largely
fluctuates.
In the second phase, some of modern humans
at the northern edge of their distribution, who
occasionally undergo loss of forests, fortunately
acquire some plain-adapted techniques andsuccessfully settle in northern plains. This causes
a population explosion of modern humans in
mid- and high-latitude areas. In mid-latitude
areas, modern humans alternately acquire and lose
forest- and plain-adapted techniques, surviving
variable vegetation conditions. Because loss andacquirement of techniques are stochastic, culturesin distant places develop more or less dependentlyb u t a s y n c h ro n o u s l y. A s a re s u l t , c u l t u r a ldiversity flourishes in mid-latitude areas, whereenvironments are most variable. While modern
humans rapidly spread over mid-latitude areas,
Neanderthals are swept out of there and become
confined within small refuge areas, such as the
Iberian Peninsula, Italian Peninsula, and Balkan
Peninsula.
In the third phase, Neanderthals remaining
in the refuges are gradually replaced by modern
humans. This phase typically lasts very long. Thisis because the refuges are permanently covered

with forests and modern humans’ innovative
ability does not help them invade. Even worse,
Mediterranean peninsulas are open only at their
northern parts, where the temperature is relatively
low and hence Neanderthals have higher viability.
Thus those peninsulas function as effective refuges
for Neanderthals, which is why it takes long formodern humans to invade them. I found that the
refuge of the Iberian Peninsula is consistently
stable and persistent compared to other refuges,
so that modern humans tend to spend most oftime in the third phase to break into this refuge.
According to this result, it is highly likely thatthe last remains of Neanderthals are found in the
Iberian Peninsula.
I also simulated a virtual world in whichclimate is stable. In this case, I found that the
replacement may or may not occur, dependingon the temperature. I.e., the replacement occurs
if the northern boundary of the forest is locatedat relatively high latitude, but it may not occur
if the boundary is located at low latitude. In thelatter case, Neanderthals permanently survive
in the Iberian Peninsula. In any case, modern
humans finally spread into plains, but this requires
far longer time than under variable climate. In
addition, the spread of modern humans into plains
does not strongly affect the population dynamics
of Neanderthals, unlike under variable climate.
Cultural diversity does not increase in any districts
without climate variation. These results suggestthat variable climate significantly affected theobserved pattern of archaeological records.

(a)

(c)

(b)

Figure 1. Simulated distributions of virtual
Neanderthals (green) and modern humans (red)in (a) the first phase, (b) the second phase, and(c) the third phase. For the definitions of the threephases, see the text.

Evolutionary Models Group B01

Figure 2. The population dynamics of virtual
Neanderthals and modern humans. The
numbers over the graph indicate the three phases(see the text).

129

A Mathematical Model of Cultural Interactions
between Modern and Archaic Humans

Wataru Nakahashi

Meiji Institute for Advanced Study of Mathematical Sciences, Meiji University, Japan

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130

O ur RNMH project is investigating the
Paleolithic cultures around the world. Now we arerecognizing that the pattern of cultural changes
in each region had been various. In Africa,
anatomically modern humans may have emerged
around 200,000 years ago, and high level cultural
traits appeared and disappeared iteratively invarious regions before 90,000 years ago but cultural
evolution speeded up and various new cultural
traits appeared after that time. In Europe, modern
humans invaded about 45,000 years ago and rapidly
spread over every region except south Iberia. The
artistic explosion of modern humans occurred soonafter their invasion. That is, certain artistic behaviors
of modern humans emerged first in Europe rather
than Africa, and some researchers consider thatinteraction with Neanderthals had influenced thesecultural innovations. In Asia, small scale invasion ofmodern humans may possibly have occurred before
74,000 years ago (inland route) and around 60,000
years ago (coastal route), and large scale invasion
occurred about 40,000-50,000 years ago. Culturalchange was not drastic but gradual in Siberia andEast Asia.
When we study these “transitional” cultures, wehave to consider the effects of interactions between
modern and archaic humans. For example, when we
study the culture of late Neanderthals, we consider
the influence of modern human culture. However,no previous mathematical models of cultural
evolution include cultural interactions. This isbecause previous models have a strong mathematical
restriction for the probability distribution ofcultural trait values each individual learns (Henrich,

2004; Kobayashi and Aoki, 2012). Therefore, I
create a new model that has no restriction for the
distribution to deal with cultural interactions.
Figure 1 describes the model. Cultural traits
are expressed as real numbers, and every matured
individual has one cultural trait (numbers in heads).
A newborn (green individual) learns cultural traits
from multiple individuals in their parental generation.
He/She explores around socially learned cultural
traits, and judges their utility following a criterion(red line) to adopt his/her mature cultural trait. He/
She sometimes blends socially learned cultural traits
to adopt the weighted sample mean, which reduces
cultural variance in the population. We consider
weak directional selection so that cultural traits withlarger value have slightly larger utility. We obtain
cultural evolutionary speed, which is defined as thegenerational change of mean cultural trait value inthe population.
First, we consider cultural evolutionary speedof single population to compare the results withprevious researches. Although previous researches
argued that larger population size and larger
number of acquaintances facilitate cultural evolution(Henrich, 2004; Kobayashi and Aoki, 2012), bothhave ignorable effect on cultural evolutionary
speed in this model provided population size islarge. This is because previous researches do notinclude blending effect, which is considered to beimportant in human cultural transmission (Boydand Richerson, 1985). Moreover, even whena fraction of population is chosen as teachers,
this effect on cultural evolutionary speed is also
ignorable provided the number of teachers is large

in Neanderthals.
Figure 4 shows the scenario of cultural evolutionin Asia. First wave invasion of modern humans
occurred when their cultural level was still low,
so that cultural interaction with archaic humans
may have affected as the rotten apple. When large
scale invasion of modern humans with higher
cultural level occurred, they may have interactedwith descendants of first wave modern humanpopulation, so that the difference of cultural levelmay not have been large. Therefore, this interaction
also affected as the rotten apple, so drastic culturalchange never occurred. In other words, intermittent
invasions of modern humans brought about thegradual cultural change in Asia.
Although these scenarios of cultural evolution
in each region may be oversimplification, it isimportant to take cultural interactions witharchaic humans into consideration when we study
transitional cultures. This mathematical model
suggests that cultural interaction can affect asboth the rotten apple and the negative exemplar. I
obtained the condition for cultural interaction toaffect as the negative exemplar. I show that whenmodern humans experienced a small interaction
with archaic humans with far lower level culture,
this cultural interaction could affect as the negative
exemplar. Although some researchers consider thatthe increase of the tension between Neanderthals
and modern humans caused the artistic explosionin Europe, this model suggests that such reason
was unnecessary provided Neanderthal culture was
low level and the cultural interaction was small.
If we consider that the increased tension caused
the explosion, it is difficult to explain why thecultural explosion occurred in art, not in weapons orsomething that was useful for competing. Moreoversince population size of Neanderthals may havebeen small compared with modern humans (Mellars
and French, 2011), the tension may not have beenstrong and the interaction may have been small.
Ignorable interbreeding between Neanderthals andmodern humans in Europe from genetic evidence(Green et al., 2010) also suggests small interactionof both species. Before the invasion of modern

Evolutionary Models Group B01

and the mean cultural trait value of teachers is thesame as that of the population, which is analogous
to Aoki et al. (2011). Furthermore, even when anewborn tends to sample individuals with similar
cultural trait, which may occur if the population is
structured, this has no effect on cultural evolutionary
speed. The width of exploring and the accuracy of
judgment have positive effect on cultural evolution.
That is, high individual learning abilities accelerate
cultural evolutionary speed, which is analogous to
Aoki et al. (2011) and Kobayashi and Aoki (2012).
Next, we study regional differences of the
Paleolithic cultural change patterns from the model.
Figure 2 shows the scenario of cultural evolution
in Africa. At a certain time modern population
with higher individual learning abilities appeared,
and cultural level of this population evolved higher.
However, cultural interaction with low ability
archaic population with low level culture began
as modern population expanded, which caused
contamination of low level culture into modern
population. This may have affected as a “rotten
apple” so that cultural level decreased until the lowlevel population went extinct. Such interaction may
have occurred several times, which entailed iterative
appearance and disappearance of high level culture
in Africa. After almost all archaic humans in Africa
went extinct, cultural evolution of modern humans
speeded up.
Figure 3 shows the scenario of cultural evolution
in Europe. Since modern humans could not invade
Europe for a long time, their cultural level may
have evolved high when they invaded Europe.
Therefore, the difference of cultural level between
Neanderthals and modern humans may have been
large. Then, Neanderthal culture may have affected
as a “negative exemplar” of how not to behave.
Because of this effect, cultural level of European
modern human population evolved higher than thatof other modern human populations that did notinteract with Neanderthals. In other words, culturalinteraction with Neanderthals may have caused
cultural explosion of European modern humans.
Moreover, cultural interaction with modern humans
may have entailed the evolution of high level culture131humans in Europe, Neanderthals had almost
no artistic materials in their culture. Therefore,
cultural difference in art was very large between
Neanderthals and modern humans, so Neanderthals
may have affected as the negative exemplar to cause
the artistic explosion of modern humans.
References
Aoki, K., Lehmann, L., Feldman, M.W.
2011 Rates of cultural change and patterns ofcultural accumulation in stochastic models of socialtransmission. Theoretical Population Biology 79: 192202.
Boyd, R., Richerson, P.J.
1985 Culture and the Evolutionary Process. Universityof Chicago Press, Chicago.

Green, R.E., Krause, J., Briggs, A.W., et al.
2010 A Draft Sequence of the Neandertal Genome.
Science 328: 710-722.
Henrich, J.
2004 Demography and cultural evolution: howadaptive cultural processes can produce maladaptive
losses-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.
Mellars, P., French, J.C.
2011 Tenfold Population Increase in Western Europe
at the Neandertal-to-Modern Human Transition.
Science 333: 623-627.

Figure 2.

Figure 1.

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132

Figure 3.

Figure 4.

An Attempt to Estimate the Functional Difference in
Haplotypes between Archaic and Modern Humans

Makoto Shimada
Fujita Health University, Japan

Background
Progress of DNA sequencing technology andbioinformatics have brought us DNA sequencesof archaic humans such as Neandertals andDenisovans. Because of post-mortem chemicalalterations such as cytosine deamination, DNA
sequences of the paleontological samples havebeen changed until the excavation. This is alimitation to estimate the functional differences
between modern and archaic humans based on
DNA sequences. To overcome this limitation, we
discovered the haplotype sequences in modern
human sequences that introgressed from archaic
to modern humans in ancient time by determining
linkage disequilibrium (LD) region using thedatabase constructed by the 1000 genomes project.

Results
We obtained SNPs that associate strongly (r2 ≥0.8) with SNPs specific to ancient haplotypes. The
obtained SNPs specific to the ancient haplotypes
contain three SNPs that are annotated to be in
the region of transcription factor binding; CDP(CUX1, Figure 1) and OCT1 found in the
intronic region of SHROOM4 gene located in theextend LD region of Xp11_hX, and CART1 thatbind to intronic region of DMD (dys44_ B006).
However, no non-synonymous SNPs were found.
This suggests that the ancient haplotype specific
SNPs affect regulation of the neighboring genes
although the protein sequences of the neighboring
genes are not changed.

Evolutionary Models Group B01

Methods
We investigated the following five sequence
elements (ancient haplotypes) that are known as
unusual haplotypes because the recent African
origin model cannot explained their divergencefrom other haplotypes in previous studies; anintergenic region on Xp11 (Xp11_hX), a regionof MCP H1 on c hromosome 8 (MCP H1_
D), a region of RPM2P4 on chromosome X(RPM2P4_AB), a region around 44-th exon of
DMD on chromosome X (dys44_B006), and ainverted region on chromosome 17 (17q21inv_
H2). To extend the genomic region of ancient
haplotype beyond that operationally determinedin the previous studies, we estimated linkage
disequilibrium (LD) regions that associate withancient haplotype specific SNPs using the data of1000 genomes project. Moreover, we annotated all
SNPs on the extended genomic regions bearing
SNPs that specific to ancient haplotypes (ancient
haplotypes).

133

Figure 1. The ancient haplotype specific SNP (rs145880787) on the transcription factor (CDP) binding site.

Evolutionary Models Group B01

134

The Down of Hierarchy: The Cognitive Ability ofArchaic Hominins

Lucas Henriques Viscardi1, Vanessa R. Paixão-Côrtes2,
Francisco Mauro Salzano2, Tábita Hünemeier2, and Maria Cátira Bortolini2
1Universidade Federal do Rio Grande – Instituto de Ciências Humanas e da Informação, Brazil

2Universidade Federal do Rio Grande do Sul – Departamento de Genética – Programa de Pós-Graduação em Genética e Biologia Molecular, Brazil

AmiGO (http://amigo.geneontology.org/cgibin/amigo/browse.cgi). Functional informationand chromosomal gene localization details wereobtained from GeneCards/GeneAlaCart database(http://www.genecards.org/). We performed allcomparisons using modern humans genomes(NCBIBuild 36/hg18), chimpanzee (2.1/ pan
Tro2 CGSC), Neanderthal and the Denisova
specimen genomes (http://genome.ucsc.edu/
cgibin/ hgBlat?org=human). All non-synonymouschanges in cognitive genes which were different
in modern humans and chimpanzees and thatwere available on Neanderthal and/or Denisova
were recovered using the USCS Table Browser.
The Grantham Score was then used to categorize
all the corresponding amino acid changes intoclasses of chemical similarity. The sites wereclassified as conservative (Grantham score 0-50),
moderately conservative (51-100), moderately
radical (101-150) and radical (>150). The AmiGO
search allowed the selection of 162 genes directly
related to cognitive process. We found 94 nonsynonymous substitutions in 52 genes, in which
the derived allele was present in humans and theancestral allele in chimp. Noteworthy 92.8% of
Neanderthal positions have the same allele foundin humans, 4.3% of the loci were heterozygote forancestral and derived alleles and 2.9% of the locishowed new variants. In the Denisova specimen
was recovered just 1 position with ancestral allele
and 1 heterozygote locus with human and chimp
substitution. From 94 substitutions, 46% wereclassified as conservative score, 38% as moderately

Evolutionary Models Group B01

D e s p i t e s h a r i n g m o s t o f t h e i r ge n e t i cbackground, humans are remarkably different
from other great apes. The most striking
difference is related to cognitive skills acquiredby Homo sapiens, and the fact that this knowledge
had been accumulated and refined throughout
its evolution as a species. Examples of these
cognitive abilities are culture, language and tool
manufacture. Within the genus Homo, thesedifferences may not be as striking. Recently withthe publication of the genomes of two archaic
Homo specimens, new discoveries have contributed
to a better understanding of the level of cognitive
development of Neanderthals species (for instancespeech ability). Humans and Neanderthals
coexisted for a long while probably directlycompeting with each other in the same ecologicalniche. Over the decades, many assumptions havebeen made ​​about the reasons for the replacementof Neanderthals by humans in Eurasia, many ofthem related to the cognitive differences between
the two species. These assumptions are basedon archaeological evidence pointing to a lower
sophistication of the development and use of tools,
as well as a less complex cultural and society in
Neanderthals. In this study we sought to evaluate
the variability in genes related to cognition inthese two species of the genus Homo as an attempt
to verify whether genetic differences in these could
be associated to Neanderthal replacement. In order
to identify genetic changes that are unique inhumans we search for genes related directly withcognitive ability using Gene Ontology database135

Evolutionary Models Group B01

136

conservative, 11% as moderately radical and 1% asradical (PDCL and CHRNA4). The PDCL geneis a putative modulator of G protein involved invarious neuronal signalizations, while CHRNA4
encodes a nicotinic acetylcholine receptor.
Noteworthy that CHRNA4 presents one change
in amino acid chain that differ from ancestralallele and Homo sapiens allele that surprisingly
reached the highest Grantham Score position(159). In human, mutations in this gene are related
to common disorders as Epilepsies Myoclonic,
Attention-deficit hyperactivity disorder, mild
mental retardation, psychiatric and neurologicaldisturbs schizophrenia and others. It is possible
to speculate whether this allele could be related
to the cognitive differences between modern andarchaic humans, notwithstanding the entire set ofdata does not point to mayor genomic differences
between the species studied. Our results suggestthat Humans, Neanderthals and Denisova
specimens are virtually equal when the molecular
aspects involved in the cognitive processes are
considered. So if there were cognitive differences
related to a less competitive Neanderthal behavior
when compared to humans, these differences haveleft no mark on genetic background. Based on
this, it seems more cautious to seek new directions
to explain the replacement of Neanderthals byhumans.

Using Numerical Models to Simulate Climate andVegetation Changes during the Last Glacial Period
Wing-Le Chan1, Ayako Abe-Ouchi1,2, Ryota O'ishi1, and Kunio Takahashi2
1Atmosphere and Ocean Research Institute, The University of Tokyo, Kashiwa, Japan
2Japan Agency for Marine-Earth Science and Technology, Yokohama, Japan

were Heinrich events (Heinrich, 1988) whichcoincided with the break-up of ice shelves in thenorthern hemisphere and the release of large
amounts of freshwater and ice rafted debris into
the North Atlantic Ocean.
Climate modellers have used numericalmodels to reconstruct the climates of OIS-3(e.g. Sepulchre et al, 2007). Paleoclimate datasuch as pollen, tree rings, ocean sediments andice cores have given us a greater insight into theclimate of the past and the way in which it has
shaped the natural environment through changes
in vegetation. In addition, they can complement
numerical models as comparisons can increase
our confidence in model simulations while
simulations themselves may give us information inregions where data are sparse. The combination ofnumerical climate models and paleoclimate datathus provides a very useful tool.
In our current work, we use numerical climate
models and incorporate proxy data to build aclearer picture of the climate during the 120,000
years so that we can better understand the its effectand that of vegetation changes on the distributionof Neanderthal and AMH population. We

Figure 1. Dansgaard-Oeschger cycles (blue)and Heinrich events (red) as inferred fromGreenland ice core data. The vertical axis
represents the isotope ratio δ18O, a measure of
the temperature which increases upward.

Climate and Geoscience Group B02

During the Paleolithic Age, hunter-gatherers
have had to adapt to changes in the naturalenvironment as a result of geographic and climatechanges. The last traces of the Neanderthals dateback to about 25,000 years ago and, at somepoint, Neanderthals were replaced completely
by Anatomically Modern Humans (AMH).
However, the role of climate in the demise of
the Neanderthals is still a subject of debate. One
hypothesis put forward is that the extinction of
the Neanderthals resulted from their inability toadapt to extreme climatic changes during oxygenisotope stage 3 (OIS-3) (Stewart, 2005). The
climate during OIS-3, which spanned roughly
65,000 to 25,000 years ago, was characterized byabrupt changes known as Dansgaard-Oeschger(D-O) events (Dansgaard et al, 1993) whichbrought about rapid warming episodes, followed bycooling over a longer period (Figure 1). It has beensuggested that these events, which appear to occur
around every 1500 years, were associated withthe changing state of the Atlantic thermohaline
circulation, weak during the cool stadials but
stronger during the mild interstadials. Precedingsome of these D-O events of the last glacial period137

have used a mid-resolution version of a coupledatmosphere-ocean general circulation model(AOGCM), called the Model for Interdisciplinary
Research on Climate (MIROC), developed at
the institutes CCSR/NIES/FRCGC in Japan(K-1 Model Developers, 2004). For time slice
experiments of the last glacial-interglacial cycle,
conditions need to be specified within themodel. These conditions are the orbital forcing,greenhouse gas concentration and ice sheet extent.
To mimic the effects of the release of freshwater
into the North Atlantic Ocean, some AOGCM
simulations include “freshwater hosing” whereby
freshwater is artificially discharged over thenorthern North Atlantic (Manabe and Stouffer,1997).
Results from these time slice experiments areapplied to a dynamic global vegetation model(DGVM), Lund-Potsdam-Jena DGVM (Sitch
et al, 2003), which is run offline to study thevegetation response to various climate scenarios(Figure 2). Climatological monthly mean valuesof precipitation and surface air temperature arecalculated over a suitable period after the climate
has reached an equilibrium state in the AOGCM.
These monthly values are fed into the DVGM
which is run with atmospheric CO2 levels specified
as before. The output data are vegetation types
represented by 10 plant functional types.
References
Banks, W.E., F. d’Errico, A.T. Peterson, M. Kageyama,
A. Sima and M.F. Sánchez-Goñi: Neanderthal
Extinction by Competitive Exclusion, PLoS ONE,Climate and Geoscience Group B02

138

Figure 2. Simple schematic diagram of a coupled
AOGCM and dynamic vegetation model.

3(12), e3972, doi:10.1371/journal.pone.0003972,
2008
Dansgaard, W., S.J. Johnsen, H.B. Clausen, D. DahlJensen, N.S. Gundestrup, C.U. Hammer, C.S.
Hvidberg, J.P. Steffensen, A.E. Sveinbjörnsdottir, J.
Jouzel, and G. Bond: Evidence for general instabilityof past climate from a 250-kyr ice-core record,
Nature, 364, 218-220, 1993.
Heinrich, H.: Origin and Consequences of Cyclic Ice
Rafting in the Northeast Atlantic Ocean during the
Past 130,000 Years, Quat. Res., 29, 142-152, 1998
K-1 Model Developers: K-1 Coupled GCM (MIROC)
Description, K-1 technical report [Eds: Hasumi, H.
and S. Emori], Center for Climate System Research,
The University of Tokyo, Japan, 34pp, 2004.
Manabe, S. and R.J. Stouffer: Coupled ocean-atmospheremodel response to freshwater input: Comparison toYounger Dryas Event, Paleoceanography, 12, 321336, 1997.
S epulchre, P., G. R amstein, M. K ageyama, M.
Vanhaeren, G. Krinner, M.-F. Sánchez-Goñi andF. d'Errico: H4 abrupt event and late Neanderthalpresence in Iberia, Earth Planet. Sci. Lett., 258, 283292, 2007.
Sitch, S., B. Smith, I.C. Prentice, A. Arneth, A. Bondeau,
W. Cramer, J.O. Kaplan, S. Levis, W. Lucht, M.T.
Sykes, K. Thonicke and S. Venevsky: Evaluation of
ecosystem dynamics, plant geography and terrestrial
carbon cycling in the LPJ dynamic global vegetation
model, Glob. Change Biol., 9, 161-185, 2003.
S t e w a r t , J. R . : T h e e c o l o g y a n d ad a p t a t i on o f
Neanderthals during the non-analogue environmentof Oxygen Isotope Stage 3, Quatern. Int., 137, 3546, 2005.

Late Middle Palaeolithic (OIS 4/3) Neanderthal Prey Exploitation and Land
Use at Dederiyeh Cave, Syria, Based on Multiple Isotopic Analyses of IntraTooth Enamel and Mortality Profiles of Wild Goat, Red Deer, and GazelleMark C. Diab and Minoru Yoneda
The University of Tokyo, JapanThe ecologist Daniel Simberloff has referred to
the final members of a species on the verge ofextinction as “the living dead” since they appear
to be healthy and active, yet lack key facets forthe species to maintain itself—to be resilient, thatinvolves social transmission, genetic drift, lackof breeding females, low recruitment (births),
increasing deaths, and negative immigration fromdwindling source areas. One interpretation may
be that Neanderthals could not recoup losses due
to an inability to “reorganize” themselves after
dynamic natural and social changes. They wereincapable of “releasing” their economic and genetic
stress to reorganize for an “exploitation” phase, thata new, more adaptable group appearing from out of
Africa could.
This research presents the first isotopic results
and mortality profiles of key herbivore preyfrom a Neanderthal site in the northern Levant,that of Dederiyeh Cave in northwest Syria.
Serial sampling from the early enamel forming
occlusal surface of the crown, to the later enamelcementum junction was conducted on a small
sample of wild goat (Capra aegagrus cf.) andred deer (Cervus elaphus). Combined results of
13
18
carbon (δ C) and oxygen (δ O), and oxygen
87
86
and strontium ( Sr/ Sr) isotopes over the 1-2.5
year period of molar enamel formation from wildgoat yield little variation reflecting local, andrestricted range habitat. This has implications
for the land use practices of Neanderthals who
occupied the region around the cave. It suggests
prior knowledge of the presence of key herbivore

Climate and Geoscience Group B02

Our closest cousins on the human evolutionaryline, the Neanderthals, sur vived for several
hundreds of thousands of years through drastic
climatic modulations that also brought withit complex ecologic al, biogeographic, andsubsistence-settlement challenges. However, muchhas been written about their extinction, and thus
apparent failure to survive, while little has beenstated about the obvious ecological and social
resilience they demonstrated through millennia ofexpansion throughout Europe and the Levant. The
question of why they became extinct still remains.
One explanation may come from a body ofconcepts defined within socio-ecological resilience
theory that attempts to explain changes in adaptive
systems that are “transforming”. Resilience theory
is based on an adaptive cycle nested in a space-timehierarchy involving stabilizing and destabilizing
effects that can cause small-scale transformations
to explode into larger-sale crises (panarchy).
In other words, unlike anatomically modern
humans, in times of socio-ecological stress, someNeanderthal groups may have lacked technological,
social, and subsistence flexibility/knowledge ina rapidly changing adaptive system and wereincapable of reorganization strategies required
during dynamic changes. On a population scale
this may have been linked to an increase in calories
that would have permitted modern humans toincrease fecundity and, thus, survivorship.
Stable isotope analysis and mortality profilesof prey species may provide palaeoecologicalsupport for extinction and replacement hypotheses.

139

Climate and Geoscience Group B02

140

prey and seasonal occurrences, and thus, potentiallyan element of complex planning and decision
making regarding the timing and schedulingof logistical or residential hunting practices.
Ethnographic evidence of wild caprid and ovid
hunting sometimes involves medium duration (afew days to one week) logistical forays involving
several hunters interacting communally. An
analogical argument from these examples can beextrapolated to hypothesize that late Levantine
Neanderthals required similar approaches to thatseen ethnographically.
Age profiles, based on time-averaged dental wear
patterns and metrics, for wild goat, red deer, andgazelle closely reflect that of southern Neanderthal
samples from Kebara cave in Israel, from a similar
period across the OIS4/3 (70,000-40,000 kya)transition. Wild goat confidence intervals on atriangular (or ternary) graph fall within an ambush
hunting strategy while red deer indicate higher
prime-age deaths, and gazelle trend toward ahigher juvenile-subadult mortality pattern. It
also suggests a consistency in hunting patterns
and prey herbivores that may have been learned
through transmission from groups close by orwithin a larger biogeographic regional boundary.
These profiles are preliminary pending a morecomprehensive investigation incorporating otheraging schemes using long bone fusion and dental
annuli. While the isotopic results are sound andprovide a stronger inferential base upon which tointerpret climatic, ecological, and biogeographic
explanations of Neanderthal herbivore prey andNeanderthal subsistence and land use at the pivotal
OIS4/3 transition, they must still be considered inconcert with potential sources of error. However,there is adequate explanator y value in thispilot study that justifies preliminary inferences
about herbivore physiology, palaeoclimate, andNeanderthal subsistence, as well as the potential
for deeper critical analysis with an augmentationof additional data in future investigations.

How Can We Apply Ecological Niche Models to
Palaeoanthropological Research?

Yasuhisa Kondo1,2 and Takashi Oguchi3
1Department of Computer Science, Tokyo Institute of Technology, Japan
2Japan Society for the Promotion of Science, Japan

3Center for Spatial Information Science, The University of Tokyo, Japan

Ecological niche modelling is a computer-based
method to simulate species’ geographical range (orecological niche) expansion or contraction based oneither real or simulated environmental conditions.
It is also applicable to palaeoanthropology andarchaeology as eco-cultural niche modelling(ECNM), because in prehistory human behaviours
were largely influenced by the environmental
impact [1]. Based on this assumption, we apply
ECNM to the spatio-temporal reconstruction of
the replacement of Neanderthals by anatomicallymodern humans (AMHs) in response to climatechange. For the practical application of ECNM,
pre- and post-processing treatments to control thedata quality and outputs are necessary. This paper
introduces techniques for preparing high resolution
palaeoclimate data and palaeovegetation zoning in
the pre-processing and for model integration in the
post-processing. Preliminary experiments are carriedout to the eco-cultural niche distribution of the
Magdalenian culture in Western Europe during theLast Glacial Maximum (LGM).

Palaeovegetation zoning
Figure 1 shows an experimental prediction of
the geographical extent of the Magdalenian nicheat 21.2 ka. The left subfigure is the prediction

Climate and Geoscience Group B02

High resolution palaeoclimate data
There is a difference in analytical scale between
the palaeoclimatic and palaeogeographic data. For
instance, the MIROC 3.2.2 atmosphere-ocean
general circulation model in the PMIP 2 protocol [2]has data points in every 2 arc-degrees. In contrast,
the resolution of the GTOPO-1 digital elevation
model (DEM) is 1 arc-minute (or 0.01667 arcdegrees). It is important for ECNM software
packages to match the pixel size of raster-based

environmental data prior to analysis. Therefore, it isnecessary to increase the resolution of the climatic
data. For this purpose, we employed the 30-arcsecond (or 0.008335 arc-degrees) climate model ofthe present day, published by the WorldClim project(http://www.worldclim.org).
The procedure of resolution increase is simple.
First, the difference between the value of the present
day and that of 21 ka was calculated for each data
point of the MIROC. For example, the differencein annual precipitation was calculated as:(1)
where ΔP is the difference, and P21k and P0k are theprecipitation at 21 ka and present (0 ka), respectively.
Then, the data points of ΔP were interpolated by aspline algorithm to create a raster surface in 1 arcminute pixel resolution. Similarly, null cells (orwater surfaces) of the WorldClim data were filled byspline interpolation and downsized to 1 arc-minute
pixel. Finally, the interpolated WorldClim data
were converted to the LGM data by subtracting thedifference ΔP.
(2)
where [Pwc_sp] is spatial matrix of the interpolated
precipitation of the WorldClim. The similar method
was applied to create the high resolution raster dataof the mean annual, warmest month (August), andthe coldest month (February) temperatures.

141

with the biome (biome model), while the rightsubfigure is the prediction without the biome (nonbiome model). Macroscopically, the two results
are similar, particular in that high probabilities
of site occurrence are predicted in the southernpart of France, Belgium, the northern half of
Switzerland, and the Cantabrian coast of northern
Spain. However, there are some differences when
we look at details: the high probability areas of
the non-biome model (right) in the west coast of
the Iberian Peninsula, the Italian Peninsula, andthe northwestern part of the Balkan Peninsula are
missing in the biome model (left). Furthermore, inthe biome model, the boundary of high and low
probability zones tends to be sharp in some parts,
which is caused by the input of the biome as acategorical value. This fact primarily indicates thata revision of the category and geospatial zoning of
biome and palaeovegetation, with reference to theplant functional type reconstructed from pollen andother palaeobotanical records, is essential to a better
application of ECNM to RNMH research. In orderto reduce the edge effect, each categorical type wasseparated and converted to a continuous value (0… 1) with fuzzy marginal buffers. Then, the sum of
the values at each cell was standardised to 1. This
treatment contributes to visualising the boundaries
of biome in a more naturalistic manner.
Model selection and integration
Among ecological niche models, GARPClimate and Geoscience Group B02

142(genetic algorithm for rule-set production) [3]and MaxEnt (maximum entropy model) [4] arecommonly applied in ECNM. Although bothmodels employ the similar inputs of location ofpalaeoanthropological sites and the environmental
dataset, these two algorithms tend to yield strikingly
different results from the same dataset and similarparameter settings. In general, the geographicalextents of the niche predicted by MaxEnt tend tobe narrower and more biased than those predicted
by GARP [5]. This tendency has also been observedin our comparative experiments. We suggest
calculating geometric mean of the standardised
GARP and MaxEnt models in order to highlightthe geographical ranges in which both models
predict high probability of species’ presence. The
contribution of each environmental variable to the
output model is also important to understand whatenvironmental factors are influential to humanbehaviour, which might have different responses toclimate change in comparison to other biological
species as a manifestation of emerging culture.
References[1] Banks W E et al. 2008. J. Archaeo. Sci. 35: 481-491.
[2] Braconnot P et al. 2007. Clim. Past 3: 261-277.
[3] Stockwell D R B. 1999. In A H Fielding (ed.)
Machine L earning Methods for Ecologic al
Applications, Kluwer, 123-144.
[4] Phillips S J et al. 2006. Ecol. Model. 190: 231-259.
[5] Peterson T A et al. 2007. Ecography 30: 550-560.

Figure 1. ECNM of Magdalenian niche range from the biome model (left) and non-biome model (right).
See also the paper by Omori and Yoneda in this volume for qualification of radiocarbon date reliability (Dat. Rank).

Bibliographic Database for “Replacement of
Neanderthals by Modern Humans”
Yuzo Marukawa1, Hirohisa Mori1, Yoshifumi Nakamura2, and Takeru Akazawa3
1Office for Virtual Resources of International Research Center for Japanese Studies, Japan
2Research Center for Informatics of Association of National Institute of Informatics, Japan
3Research Institute of Kochi University of Technology, Japan

Creation of bibliographic data
We have been constructing a bibliographic
database for “Replacement of Neanderthals byModern Humans” covering reports on relics
and remains of relevance to the Replacement of
Neanderthals by Modern Humans project. By theend of 2011, we had successfully digitized a total of
2,713 pieces of bibliographic materials, including
full-text conversion of 60,613 pages.
Subsequently, we created metadata for each ofthese bibliographical materials, adding remain
name, location, era, keywords and other pertinent
facts, as well as standard bibliographic informationsuch as title, author, publication year, journal name,volume number, and pages. We initially createdthe catalog data in Excel files and, after importingthem into the bibliographic search system, edited
the data directly from Web browsers.

Figure 1. Search screen of the “Replacement of
Neanderthals by Modern Humans” bibliographic
database.

Climate and Geoscience Group B02

Bibliographic search system
We constructed a dedicated bibliographic
search system for the purpose of searching forand browsing bibliographical materials utilizing
the aforementioned digitized literature metadata.
This system stores the bibliographic data and iscomposed of a search environment for users (frontend) and a data editing environment (back end).
The front end allows searching by free text,
author, publisher, and publication year. For authors,in particular, the system is equipped with a socalled suggest function, which makes an indexedauthor list from the database and displays likely
author candidates in response to inputs from users.

In addition, search results are displayed in order ofauthor names. If there are multiple author names
matching the search, the results are sorted by the
first author name and displayed in the ascendingorder of the number of authors and then the
publication year. The list of search results shows
bibliographic information as well as links to fulltext PDF files (Figure 1). Users can click on thelinks to immediately display the full-text PDF filesof the relevant literature.
The back end has an interface that allows editing
contents of the database. Users with editing rights
are able to use internet browsers to list and edit
data. We use this back end to correct typographicalerrors, characters omitted in the text conversion
process etc., and add related information. Since itallows searching for target literature and working
on the data while displaying full-text PDF files of143

the target literature, we are able to edit the data
more efficiently than by working with Excel files,
which we did at first.
Coordination with external databases using
associative technology
Creating the bibliographical material database
opened up the possibility of linking with external
databases. We thus made use of the Associative
Retrieval System “IMAGINE” Book Search tobuild a service for looking up items pertinent totarget literature from existing fossil databases.
Moreover, taking advantage of the literaturebrowsing system “eReading” (Figure 2), weconstructed a service that allows looking up item
names in fossil databases and encyclopedias whilebrowsing literature PDF files. This service utilizes
functionality from the “Ebiki” system that allows
attaching annotations to images.
Coordination with these systems allows us todirectly add external bibliographic data to ourbibliographic database. To further facilitate theintegration, we employed a data format that is
relatively easy to handle in our bibliographic
database and allows partial update of the data
simply by specifying an ID for the data.
Utilization of the bibliographic database andeffort toward construction of remains database
Our purposes for constructing the bibliographic

Climate and Geoscience Group B02

144

database are not limited to searching for literature
itself. We have another purpose as well; fulltext data of bibliographic materials contain large
volumes of descriptions of facts and figures, andwe want to be able to browse such information
systematically as well. In case of remains inparticular, the actual sites may be lost duringarchaeological excavation, and descriptions ofremains in bibliographic materials thus carrysignificant weight.
Towards this end, as part of our study ofapplications of the bibliographic database, we areworking on design and development of systems
used to organize and deliver information focusing
on remains. Specifically, we created test data anddeveloped a prototype database of remains, byreferring to the recent report on the Dederiyeh
Cave (Akazawa and Muhesen eds. 2003). In
the development, we focused on monographfigures and photos in the paper and attempted todesign the system to extract and format as muchinformation on remains as possible from thebibliographic materials. For example, in order todirectly link captions to figures, we made use of
the aforementioned functionality for attachingannotations to images and incorporated latitude/
longitude data of shooting locations, shooting
directions and other elements into photos. We aim
to upgrade and improve the delivery environmentas part of our future development of the system
and achieve a service environment that allows
browsing minute information on remains.
Reference
Akazawa, T. and S. Muhesen (eds.)
2003 Neanderthal Burials - Excavations of theDederiyeh Cave, Afrin, Syria. Auckland : KW
Publications Ltd.

Figure 2. Linkage between body text and pertinent
items via “eReading”.

Plant Creating a Common Chronology for Levant
Pollen Records Based on Ice Core Synchronization

Stephen Obrochta, Yusuke Yokoyama, and Hodaka Kawahata
Atmosphere and Ocean Research Insitute, The University of Tokyo, Japan

determined though the summing of annuallydeposited snow layers with a similar accuracy asradiocarbon dating. For example at 50,000 years,
the Maximum Counting Error of the NorthGRIP (NGRIP) ice core is approximately ±1,000 years (1σ), which is roughly comparable
to the uncertainty in the latest radiocarbon
calibration (IntCal09) prior to the addition of
measurement error. Therefore, the combinationof high precipitation rates in Greenland with thelarge spatial extent of the synchronous temperaturechanges provides a powerful tool for assigning agesthat would otherwise be unknown.
However, there have been significant revisions
to the Greenland ice core chronology over the past
15 years, and many of the published age models
of both marine and lacustrine sediment cores are
not compatible. The largest offsets, up to 4,000
years, are between the early-glacial sections of the
GISP2 and NGRIP ice cores, which is the exact
interval in which radiocarbon is no longer of use.
Up until 2005, the Meese-Sowers 1994 (MS94)
layer-counted chronology for the GISP2 ice core
was the most widely used correlation target.
Since that time, very high density sampling of
the newer, virtually complete NGRIP ice core
indicates that GISP2 likely suffers from excessive
layer thinning, and the NGRIP GICC05 layercounted chronology, extended with the ss09sea
flow model, has subsequently replaced GISP2
as the preferred chronology. Comparing older,
GISP2-aged data with newly published datarequires updating the former to the GICC05

Climate and Geoscience Group B02

The early portion of the last glaciation isbeyond the range of reliable radiocarbon dating.
This is problematic for assigning ages to an
important period in human history that includes
the migration of modern humans out of Africa
and their eventual replacement of Neanderthals.
In addition to the assignment of absolute age,
this also complicates understanding the relative
deposition timing of stratigraphic layers fromdistant sites.
An alternate dating method is the comparisonof environmental proxy records, obtained
from the area of interest, to the Greenland ice
cores air temperature records. Multiple largeamplitude, abrupt warmings, commonly known as
Dansgaard-Oeschger (D-O) Events, punctuatedthe last glaciation. A large body of work indicatesthat these interstadial events are expressed ascontemporaneous environmental fluctuations overa wide spatial area, centered on the North Atlantic
Ocean and adjacent continental regions, with farreaching connections to more distal areas.
North Atlantic sea surface temperature proxyrecords obtained from marine sediment cores often
exhibit a warming and cooling pattern comparable
to Greenland. Similarly, in continental regions,pollen-based environmental reconstructions
indicate the return of relatively warm and humid
conditions during Greenland interstadials. Within
the range and resolution of radiocarbon dating,these fluctuations occur simultaneously with
Greenland air temperature changes.
The age of the ice in Greenland cores may be

145model-extended chronology.
Here we present an automated mathematical
function for doing so using either the Matlab
or Octave (free, open source version of Matlab)
high-level numerical computational languages.
The function is based on synchronization of theice cores and allows for specifying the age of theoldest radiocarbon data as a starting point for
converting any input series from the MS94 to the
GICC05-extended chronology. The output ages
are relative to the number of calendar years before
AD 2000 (i.e., kiloyears b2k). Dates outside of
the GISP2-to-NGRIP conversion range are notadjusted except for adjust to the b2k scale. This
will facilitate immediate comparison of older andnewer data and will be the basis of a consistentchronology for pollen records obtained from theLevant region.

Climate and Geoscience Group B02

146

Population Dynamics and Spatial Distribution Inferredfrom Radiocarbon Meta Analysis

Takayuki Omori and Minoru Yoneda
The University Museum, The University of Tokyo, Japan

resolution is required.
The reliability assessment of radiocarbon dates
follows a decision tree, which is constructed frompurely technical standpoint. For the decision tree,
we set up mainly four evaluation criteria: oldness ofradiocarbon age, margin of 1σerror, sample material
type and technical comment on the publication. The
data reliability is rated on a scale of one to four. The
reliability score 1 is ruled unfit to apply the meta
analysis, and rejected in this study.
For radiocarbon age calibration and meta analysis,
we use OxCal 4.1 and 4.2 beta program (Bronk
Ramsey, 2009 and 2012). All of the accepted data
(score 2 to 4) was calibrated by Intcal09 calibration
curve (Reimer et al., 2009), and then carried out
radiocarbon data analysis. The calibrated dates wereclassified into lithic industries as population groups,
and the temporal frequency distributions were
calculated from the data groups using radiocarbon
Bayesian analysis.
Figure 1 illustrates the temporal distributionof Mousterian, Aurignacian and Gravetian lithic
industries throughout the European region.
Patterns of the distributions were altered by theassessment evaluation, especially in later period ofeach industry. In the Mousterian industry, which is
Neanderthals lithic culture, there are no substantial
fluctuations in the duration, and the frequency has
reduced gradually over time. On the other hand,
there are large changes in the distributions of the
Aurignacian and Gravetian as Modern humancultural groups around 35,000 BP and 30,000 BP.
The spatial distribution from the evaluated data is

Climate and Geoscience Group B02

Meta anal y sis of r adioc ar bon databaseprovides information about population
dynamics independently of archaeological andanthropological approaches. Assuming the datablesample, such as charcoal, human skeleton andanimal bone, corresponds to evidence of humanactivity, the number of reported radiocarbon
dates can be interpreted as qualitative population
dynamics. Several studies have been reported
on the radiocarbon meta analysis for the Upper
Paleolithic in Europe, and the radiocarbon-based
chronology as well as its frequency distribution
have been discussed to understand the replacement.
In this study, we evaluate the publishedradiocarbon dates and reconstruct the chronologyof the Upper Paleolithic in Levant and Europe.
Radiocarbon dates and archaeological information
were taken from our own chronological database“NeandatDB” and PACEA geo-referenced
radiocarbon database (d'Errico et al., 2011).
In addition we attempt to estimate population
dynamics and its special distribution.
Generally almost all radiometric databases
include a number of problematic dates fromthe viewpoint of technical confidence in themeasurement, adequacy of sample material andso on. Most of them are reported during an early
period of radiocarbon research. Even though someunreliable dates incorporated in database, the meta
analysis would be able to exhibit rough outline of
the chronology if mass data was used. However,when comparing the obtained chronology andabrupt paleoclimate events, more high temporal

147

shown in Figure 2. Although the distribution of the
Neanderthals has been contracting over time, thedistribution area of Modern humans seems to nochange from the 40,000 BP despite some fluctuationsin the temporal distribution. In this paper, we discussthese time and spatial distributions in detail, andestimate the population dynamics based on the result.

Figure 1. Temporal frequency distributions of theMousterian, Aurignacian and Gravettian lithic
industry. Each solid and dashed line was derived fromonly the acceptable radiocarbon data and all the data inthe database respectively.

Climate and Geoscience Group B02

148

Figure 2. Special distribution of Neanderthal andmodern human groups based on the evaluatedradiocarbon data. The size of plotted circle represents
the probability of the existence of population groups.
Red and blue circle belongs to the Mousterian industry
as Neanderthals and the Aurignacian and Gravettianindustry as Modern humans respectively.

PaleoGeo: A Web GIS for Distributing Informationfrom Paleoenvironmental LiteratureSong Wonsuh1, Yasuhisa Kondo2,3, and Takashi Oguchi1
1CSIS, The University of Tokyo, Japan

2Department of Computer Science, Tokyo Institute of Technology, Japan
3Japan Society for the Promotion of Science, Japan

There is a growing interest in the paleoenvironment,
and the number of relevant research papers has been
growing rapidly. Paleoenvironmental studies covervarious fields such as paleohydrology, geomorphology,
paleooceanology, paleobiology, paleoclimatology,
and chronology. It is hard to collect and compile
these enormous data by an individual researcher.
For the multidisciplinary project, “Replacementof Neanderthals by Modern Humans (hereafter
RNMH)”, we have been compiling portal data byusing a web-based geographical information system(Web-GIS). We call this system “PaleoGeo”. It helps

an individual researcher to search paleoenvironmental
information efficiently.
The PHEIMS (Paleo-Hydrology and Environment
Internet Map Server), a precursor of the PaleoGeo,
was developed in the late 1990s by one of the coauthors (T. O.), using ESRI ArcView IMS as a main
server software package. It contained informationfrom papers in international journals of earth andQuaternary sciences published during the mid-1990s
to 2002. It was viewed up to 3,000 times annually;
however, its updating ceased because of the end of
the relevant project.

Climate and Geoscience Group B02

Figure 1. Interface of the PaleoGeo. Yellow pins show sites with information such as location (latitude
and longitude), bibliographic data, keywords, and period. Each record is linked to the publisher’s abstract
website using a permanent link (DOI).

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The PaleoGeo is being developed to revive andupdate the PHEIMS. We improved the userinterface and data quantity (papers up to year 2011).
The new system uses ESRI ArcGIS Server 10 and to
reduce redundancy of the data, a relational databasemanagement system (RDBMS) is applied.
The data consist of the journal name, information
about each paper (authors, title, volume, year,
and page numbers), site location (country name,longitude, and latitude), theme, subtheme, keywords,
DOI (Digital Object Identifier), and period (era).
DOI enables users to see publisher’s abstract pages in
one click. Bibliographic information is also available
at some existing web-based search engines such as
the ISI Web of knowledge. However, location data
are indispensable in paleoenvironmental studies. The
PaleoGeo shows information with a map (Figure 1),
which is the most distinguishable advantage of thisdatabase system.
We have collected information from almost
three thousand articles of 13 journals regarding
paleoenvironmental research (i.e., Boreas, Catena,
Climatic Change, Earth Surface Processes andLandforms, Geomorphology, Journal of Quaternary

Climate and Geoscience Group B02

150

Figure 2. Part of original data after a search query.

Science, Palaeogeography, Palaeoclimatology, andPalaeoecology, Quaternary International, Quaternary
Research, Quaternary Science Reviews, The
Holocene, and The Journal of Geology). The articles
were classified into six themes (paleohydrology, earth
surface processes and materials, paleooceanology,
paleobiology, palaeoclimatology, and chronology) and19 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). The original data canbe viewed as a search result (Figure 2). Until now,
information of 6,881 sites has been collected and thenumber is being increased.
Although the PaleoGeo seems to be a useful tool,
it has room for improvement; for example, expanding
a window showing search results, and improving themap print function. We will fix such problems assoon as possible. In the future, this database system
will be connected to the holistic bibliography catalogof the National Institute of Information (NII), Japan.
This will be another way for the effective use of thesystem.

Statistical Interpolation of Missing Parts in Fossil
CraniaHideki Amano1, Yusuke Morita1, Hiroyasu Nagano1, Osamu Kondo2,
Hiromasa Suzuki3, Masato Nakatsukasa4, and Naomichi Ogihara1
1Department of Mechanical Engineering, Faculty of Science and Technology, Keio University, Japan
2Graduate School of Science, The University of Tokyo, Japan

3Research Center for Advanced Science and Technology, The University of Tokyo, Japan
4Department of Zoology, Graduate School of Science, Kyoto University, Japanand other remaining coordinates as independent
variables. Let Δpi (i=1..N) be the (3×1) position
vector of the ith missing landmark from theconsensus cranium and Δx be the (3(110-N)×1)
vector representing the deviations of the remaining
landmark coordinates from the consensus (N is thenumber of the missing coordinates). The regression
equations can be written as:						
(1)
where Ai is the (3×3(110-N)+1) matrix of regression
coefficients for Δpi. To solve for Ai, we exploitedcorrespondence between Δpi and Δx in the complete
specimens. Specifically we solved the following
simultaneous equations for Ai.
			(2)
where Δpi,j and Δxj (j=1..S ) be the Δpi and Δx of
the jth complete specimen, respectively, and S is thenumber of the reference complete specimen. As
the number of equation is less than the number of
unknown quantity, the solution is underspecified.
Hence we used the Moore-Penrose pseudo-inverse
matrix to obtain the solution. In order to examine
the efficacy of the proposed interpolation method,
we virtually created crania with missing portion.
Figure 1A shows the virtually generated crania with
5 (top row) and 20 (bottom row) missing landmarks,
respectively. The missing landmarks in the crania
were then estimated by using Equation (1) and theestimated coordinates of the missing landmarks werecompared with their true values.

Skull and Brain Group C01

During fossilization, crania are often fractured andall the component fragments are rarely recovered.
To establish anatomically accurate interpolation of
missing parts of fossil crania is of crucial importancefor correct estimation of brain morphology basedon cranial shape information. In the present study,
therefore, we attempted to establish a method tostatistically interpolate missing coordinates of crania
based on reference database of cranial morphology.
To construct reference database of human cranial
shape, we used a total of 22 crania (9 female and 13
male crania) from the modern Japanese population
housed at Kyoto University. Each cranium wasscanned using a CT scanner and the 3D surface modelof the cranium was generated. On the surface of eachcranium, 17 anatomical landmarks were digitized.
Furthermore, the midsagittal, superior nuchal, andtempral curves were approximated using a seventhorder Bezier curve and equally spaced points alongeach of the curves were also defined as landmarks.
To capture the morphology of the entire cranial
vault, 39 shortest path connecting pairs of landmarks
were calculated and equally spaced points alongeach of the paths were also defined as landmarks.
Therefore, a total of 110 landmarks were acquired foreach specimen to describe the biological shape. The
landmark coordinates were then normalized by thecentroid size and registered using the Generalized
Procrustes Analysis.
Based on a sample of complete specimens as
reference data, multivariate regressions are calculatedwith the missing coordinates as dependent variables

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Figure 1B compares the estimated positions of themissing landmarks with those of the original cranium.
Here the neurocranial shape is represented by thewireframe connecting the landmarks. As illustrated,the estimated positions of the missing landmarks
are located reasonably close to the corresponding
landmarks on the original cranium. The mean
prediction errors (± standard deviation) were 2.6 ±1.6 mm and 2.9 ± 2.3 mm for the crania with small
and large missing portions, respectively. Althoughintroduction of adequate constraints seem necessary
since solving Equation (2) is an ill-posed problem,the present interpolation framework may contributeto precise estimation of missing parts in fossil crania.

Figure 1. (A) Virtually generated crania with missing
portion. (B) Comparisons of the estimated positions of themissing landmarks with those of the original cranium.

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CT Image Segmentation Using FEM with Optimized
Boundary Condition

Hiroyuki Hishida1, Hiromasa Suzuki1, Takashi Michikawa1,
Yutaka Ohtake1, Satoshi Oota2, Naomichi Ogihara3, and Osamu Kondo1
1The University of Tokyo, Japan
2RIKEN, Japan

3Keio University, Japan

based on the use of structural analysis for CT
image segmentation. However, it was necessary
with this approach to set boundary conditions
manually in structural analysis. Thus trial and error
is required to apply the method.
In this paper, we expand our previous work
to set parameters and loading conditions in asemiautomatic way. We set regions of interest(ROIs) as candidates for weak areas, which
include correct and erroneous segmentation
targets. Assuming strain as an objective function,
we calculate boundary conditions for each ROI
using a gradient ascent method. In other words,
we calculate strain for all ROIs. Then, we choose
the ROI with the highest strain as the correct
candidate for a weak area, and these conditions arethen used to segment the image.
The main advantage of the proposed approach is
its reliability, as the structural analysis it involves
means that appropriate object segmentationin structurally weak areas is guaranteed. As
this method is designed to segment joints ofa skeleton and thus prefers such objects withshapes of relatively high aspect ratio around thesegmentation boundaries. The originality of thisresearch lies in its introduction of the analogy ofmechanical destruction to image processing and its
optimization of boundary conditions in the finite
element method (FEM).
Green area in the top and left middle are ROI
and the red arrows show some representatives ofloading forces. Images in the right and middle

Skull and Brain Group C01

We propose a CT image segmentation method
using structural analysis. Motivation of ourresearch is from the area of fossil reconstruction
to decompose assembled fossil of skeletons andcrania to their fragments. X-ray CT is used toscan an assembly then its image is manuallysegmented into pieces of the bones. Considering
the number of fragments involved, it is desirable
to make this segmentation procedure automatic. A
problem specific to this segmentation is to separate
fragments where their gap are not necessarily
clear. Despite the numerous image segmentation
methods, none of them matches our objectives.
Accordingly, we previously proposed a method
to segment CT images using structural analysis.
The technique is based on the assumption thatthe interference area (joint) between components(bones) is structurally weak. We compute strain,
which tends to be large in structurally weak areasand segment the image in the region of high strain.
In previous work, we used commercial software
VOXELCON, which is an image-based structural
analysis system. In the approach examined, we
set physical properties for every pixel and createa stiffness matrix. Then, we calculated the vonMises strain of every pixel and removed the one
with the highest strain value, assuming it will be
broken. We iterated strain calculation and removal
until the input image is decomposed into multiple
fragments. The method could be used to segment
low-contrast CT images - a task that was difficult
with conventional methods. This technique was153show distribution of von Mises strain andred pixels indicate high value. Using analogyof destruction we repeat removing these red
pixels until the object gets separated along theinterference region. In the bottom, the result ofsegmentation is shown where the removed pixels
are salvaged to be included in the segments.

Figure 1. Flow chart of the proposed segmentation algorithm.

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154

Cerebellar Size Estimation from Endocranial
Measurements

Daisuke Kubo1 and Hiroki C. Tanabe2
1Graduate School of Science, The University of Tokyo, Japan

2Graduate School of Environmental Studies, Nagoya University, Japanestimated these by using the regression equation
based on PCF volume of a mixed hominoid species
sample (MRI data of 4 hylobatids, 13 great apes,
and 17 modern humans) (Weaver, 2001), and she
found that the Neanderthal specimens had smaller
cerebellar volumes, relative to their whole brain size,
than those of the living human sample (Weaver,
2001, 2005). She justified her estimation by thehigh correlation between the PCF and cerebellar
volumes from MRI data (r2 = 0.89) (Weaver, 2001,
2005). However, the high correlation was probably
due to a large range of variation in size fromgibbons to humans, and it does not mean that the
cerebellar volume estimates of the fossil specimens
are accurate. The first author and co-workers reexamined the Weaver’s dataset and found that
correlation within the human sample (n=17) isactually weak (r2 = 0.16) and not significant (Kuboet al., 2011). Thus the suggested difference of thecerebellar volume between the Neanderthals andmodern humans remains to be tested.
In order to develop estimation methods of thecerebellar volume and also clarify the limitation,
we examine the correlation between the PCF andcerebellar volumes using MRI data of Japanese
subjects (Figures 2 and 3). We found that the twovolumes are more highly correlated in our dataset(Figure 4) than expected from Weaver’s MRI dataof the human sample. The PCF volume in our
definition do not include the volume of the anterior
part of the PCF region where the brain stem islocated (Figures 2 and 3), and this could partly
contribute to the high correlation with the cerebellar

Skull and Brain Group C01

There is a broad consensus that higher cognitive
functions in modern humans, including language,
are supported by the well-developed neuralbases, especially those of the cerebral region(e.g. Beaumont, 2008). On the other hand, anincreasing number of neuroanatomical, clinical, andfunctional neuroimaging studies suggest that the
cerebellum also contributes to cognitive functions(Murdoch, 2010; Stoodley and Schmahmann,
2010). Therefore, reliable data of cerebellar volume
estimates of extinct hominins can be an importantsource in understanding when and how the neuralbases related to higher cognitive functions evolved.
Especially, whether Neanderthals were significantly
different or not from modern humans in the
cerebellar volume is an intriguing question because
these two species differed in behavioral and cultural
aspects (e.g. Klein, 2009) although they were largely
comparable in the whole brain size (estimated fromthe endocranial volume) (Schoenemann, 2006).
The cerebellar volume of fossil hominins might
be estimated from the posterior cranial fossa (PCF)(Figure 1). The PCF is a posteroinferior region of
the endocranial cavity and a bowl-shaped depressionthat houses a substantial volume of the cerebellum.
However, the relationship between the PCF and the
cerebellum is not so simple; an anterior space of the
PCF are occupied by the brain stem, while an upper
portion of the cerebellum protrudes from the upper
border of the PCF.
Cerebellar volume estimates of some fossil
hominin specimens have been reported by thepioneering work of Weaver (2001, 2005). She

155

volume. At any rate, the fact that a high correlation
was confirmed in a modern human population could
offer promising prospects for its application towards
the other modern human and possibly Neanderthal
examples.

References
Beaumont J.G. (2008) Introduction to Neuropsychology,
2nd edition. The Guilford Press, New York.
Klein R.G. (2009) The Human Career: Human Biologicaland Cultural Origins, 3rd edition. The University of
Chicago Press, Chicago.
Kubo D., Kono R.T., and Suwa G. (2011) A micro-CT
based study of the endocranial morphology of theMinatogawa I cranium. Anthropol. Sci., 119: 123-135.

Murdoch B.E. (2010) The cerebellum and language:
historical perspective and review. Cortex, 46: 858-868.
Schoenemann P.T. (2006) Evolution of the size andfunctional areas of the human brain. Annu. Rev.
Anthropol., 35: 379-406.
Stoodley C.J. and Schmahmann J.D. (2010) Evidence for
topographic organization in the cerebellum of motor
control versus cognitive and affective processing.
Cortex, 46: 831-844.
Weaver A.G.H. (2001) The Cerebellum and Cognitive
Evolution in Pliocene and Pleistocene Hominids.
Dissertation, University of New Mexico, Albuquerque,
NM.
Weaver A.H. (2005) Reciprocal evolution of thecerebellum and neocortex in fossil humans. Proc. Natl.
Acad. Sci. USA, 102: 3576-3580.

Figure 1. Superior view of thelower half of an endocranial cavity,
rendered from CT data. The PCF isthe depression area surrounded by agreen line.

Figure 2. The polygon surface
model of the PCF region. An PCF
region is first segmented from MR
volume data, and then converted intoa polygon surface model. The volume
was measured after trimming by theanterior and superior planes defined byanatomical landmarks.

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Figure 4. Bivariate plot of the PCF
and cerebellar volumes with reduced
major axis (a preliminary result
based on a small number of sample).

Figure 3. Superimposed image of the
PCF region and cerebellum.

Transferring Semi-Landmarks on Surfaces

Masaki Moriguchi1, Hiromasa Suzuki2, Takashi Michikawa2,
Naomichi Ogihara3, and Osamu Kondo4
1Department of Information and System Engineering, Chuo University, Japan

2Research Center for Advanced Science and Technology, The University of Tokyo, Japan
3Department of Mechanical Engineering, Keio University, Japan

4Department of Biological Science, The University of Tokyo, Japan

MethodWe assign an Affine transformation to each vertexof the mesh as in [1] and compute it by minimizing
the bending energy. Since the landmarks PA on A

corresponds to the landmarks PB on B, we set hardconstraints: PA is transformed to PB. This constrained
optimization can be solved by the Lagrange
multiplier method and it reduces to the linear system
as shown in [5].
Letbe the Affine transformation assigned to the i-th
vertex of the surface mesh A. Then, for x-coordinates,
we solve the following linear system:

where L is a Laplacian matrix, . denotes the
Kronecker product, I4 is the identity matrix of size
4, ax is a variable that corresponds to the Affine
transformations, C ax = bx is the set of the hardconstraints, and λx is the Lagrange multipliers.
Specifically,2ax =. LT L2
corresponds to the bi-Laplacian system, and the i-th
row of C ax = bx corresponds to the constraint that piA
is transformed into pib (i = 1…k). The linear systemis large and sparse, and its size is (4n + k)×(4n + k),
where n is the number of the vertices of the surface
mesh A and k is the number of the landmarks. We
have similar linear systems for y- and z-coordinates.
By solving these three linear systems, the Affine
transformations of the vertices are obtained.
Note that, since the coefficient matrices of theSkull and Brain Group C01

Introduction
Estimating the positions of semi-landmarks isimportant in analyzing the fossil crania [4]. While
we can locate anatomical landmarks reliably, locatingsemi-landmarks consistently is a difficult task. Thisis because semi-landmarks are generally distributed
around featureless regions and we cannot locate themusing local shape descriptors.
To estimate the positions of semi-landmarks, weconsider the problem of semi-landmark transfer inthis research: given a surface mesh A (with landmarks
PA and semi-landmarks QA) and a surface mesh B(with landmarks PB), transfer the semi-landmarksof A onto B. In other words, compute the positionsof semi-landmarks QB of B using PA, QA and PB. We
assume that PA = {piA}i =k1 and PB = {piB}i =k1 have thesame size, and that each landmark piA on A has acorresponding landmark piB on B (i = 1…k).
Sliding semi-landmarks [3] is the standard
method in semi-landmark transfer. In this method,semi-landmarks are transferred so as to minimize
the bending energy of the thin-plate spline warp
[2] between PA ∪QA and PBA ∪QB. We also use anenergy minimization approach, but we minimize thebending energy over the surface instead of over thewhole 3D space (which is the case of sliding semilandmarks).

157

linear system for y- and z-coordinates are the sameas that of the linear system for x-coordinates, we
can reuse the decomposition results to save thecomputational time of the linear solves.
References[1] Allen, B., Curless, B., and Popovic, Z. The space ofhuman body shapes: reconstruction and parameterizationfrom range scans. In Proceedings of SIGGRAPH, 587–594,
2003.
[2] Bookstein, F. L. Principal warps: thin-plate splines andthe decomposition of deformations. IEEE Transactions on
Pattern Analysis and Machine Intelligence, 11(6): 567–585,
1989.
[3] Bookstein, F. L. Landmark methods for forms without
landmarks: morphometrics of group differences in outline
shape. Medical Image Analysis, 1(3): 225–243, 1997.
[4] Gunz, P., Mitteroecker, P., Neubauer, S., Weber,
G. W., and Bookstein, F. L. Principles for the virtual
reconstruction of hominin crania. Journal of Human
Evolution, 57(1): 48–62, 2009.
[5] Welch, W. and Witkin, A. The space of human body
shapes: reconstruction and parameterization from range
scans. In Proceedings of SIGGRAPH, 157–166, 1992.

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158

A Geometric Morphometric Study of Neurocranial
Shape Variations in the Crania of Modern Japanese

Yusuke Morita1, Hideki Amano1, Masato Nakatsukasa2, Osamu Kondo3
and Naomichi Ogihara1
1Department of Mechanical Engineering, Faculty of Science and Technology, Keio University, Japan
2Department of Zoology, Graduate School of Science, Kyoto University, Japan
3Graduate School of Science, The University of Tokyo, Japanpositions of the sliding landmarks on each of thespecimen. The coordinates of the landmarks werethen analyzed using Morphologika (http://hyms.
fme.googlepages.com/resources). The landmark
coordinates were normalized by centroid size
and registered using the Generalized Procrustes
Analysis. Principal components (PCs) of shapevariations among the specimens were thencalculated using the variance-covariance matrix of
the Procrustes residuals of all crania.
Figure 1B shows the variations in the 3D shapealong PC1 by warping the neurocranial shape
represented by the wireframe connecting thelandmarks. The PC1 accounted for 24.1% of thetotal variance. We observed that with increasing
PC1, a relative contraction of the cranial lengthand relative elongation of the cranial breadthwere observed. Particulary, the forehead andoccipital region had receded and the nuchal lines
were located relatively located more inferiorly
with an increase in PC1. We demonstrated thatthe most predominant shape variability in the
cranial vault in the modern Japanese populationis the brachycephalic/dolichocephalic tendency.
We will use this detailed quantification methodof the human neurocranial shape to constructa morphological database of human cranial
shape, which is indispensable for mathematicalreconstruction of fossil crania.

Skull and Brain Group C01

Geometric morphometrics have been widelyused for the analyses of morphological variabilities
in human crania. However, mor phologicalvariability in the neurocranial shape has not been
well documented because only a few definable
landmarks exist on the human cranial vault. In
the present study, we analyzed the morphologicalvariability of the neurocranial shape in the modern
Japanese population using the sliding semilandmark method.
We used a total of 60 crania (27 female and 33
male crania) from the modern Japanese population
housed at Kyoto University. Each cranium wasscanned using a CT scanner and the 3D surface
model of the cranium was generated. Template
landmark configuration that will be projected ontoand slid along each of the samples was preparedbased on the shortest paths connecting pairs ofanatomical landmarks. Specifically, conventional
anatomical landmarks and equally spaced points
along the superior nuchal curve and temporal
curve are defined as non-sliding landmarks.
These non-sliding landmarks as well as equally
spaced points along the midsagittal curve and thefrontal torus were used to calculate the shortest
paths as shown in Figure 1A and the equally
spaced points along them were designated as
sliding semi-landmarks. Therefore, the template
landmark configuration consisted of a total of 121
sliding and non-sliding landmarks. In the present
study, Templand in the EVAN Toolkit (http://
www.evan-society.org) is used to calculate the159Figure 1. (A) Template landmark configuration. (B) The
variation in neurocranial shape represented by PC1.

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160

Motion Analysis for Stone-Knapping of the Skilled
Levallois Technique

Yukinobu Hoshino1, Naoki Miura2, and Kenji Nagai3
1School of Engineering, Kochi University of Technology, Japan
2Faculty of Engineering, Tohoku Institute of Technology, Japan

3Department of Historic Heritage, Tohoku University of Art & Design, Japanaction getting Levallois flakes. In this project, learning
process is main topic. Learning (education) is for thesurvival of mankind. Archaeological material is the
only evidence that indicates the learning behaviorof prehistoric times. We collect and analyze thearchaeological materials and want to find out what's
the difference between human beings and the New
Learning of archaic humans. Levallois techniqueand skill is one kind of the archaeological materials.
Modern humans begins with a cognitive behavior in
specific, they obtained this skill during short time at
250,000 years to 6 million years. We are consideringthat those are from the evolution and a groupproduction by mixing the group learning and theindividual learning.
Two process of Levallois technique for the motion
analysis
It is important to analyze how skill is acquired,in order to understand human's learning ability. In
this study, we analyzed a difference of kinematic
information of hand motion between each procedure
to construct a stone-tool by the Levallois techniquefrom a 3D motion data measured from a skilled
subject. For those analyses, we consider 2 levelprocesses about Levallois technique, See Figure 1. We
catalogs two process data classes. The first process is abeat knapping to make a form called a stone testudinalstyle. Second process is a strong knaps to flake off apiece. So, ① is in the process of forming the testudinalstyle by finely knap. The core stone is deformed visually
by a beat knapping, so the former is possible to checkvisually and acquired by social imitation. The knapper

Neuroscience Group C02

Introduction
An investigation of skilled behavior is important toclarify a cognitive mechanism of learning process. In
this research, we measured 3D motions of stone-tool
production based on the recurrent Levallois technique
by a highly skilled subject. And, a motion analysisof measured kinematic information was performed.
To combine the analysis based on the experimentalarchaeology, it will be available to extract body motions
which is represented the skilled behavior and to
estimate a force necessary for processing of Levallois
points (LP) and Levallois flakes (LF). We would liketo report the analysis results about expert technology
of Levallois. In January 2011, we acquired the motionof the chipped stone tool using the motion capturesystem. In this experiment, the subject is expert masterof Levallois technique. The subject formed code's
edges at one end. After that the subject pick up atarget flake by flaking off pieces around the outlineof the intended flake. The subject creates those flakes
by strike hammer-stone repeatedly. On using the
motion capture system, three tracers are put aroundthe target subject. The tracers took 3D motion data
subject and hummer stone. Those data has included
the acquisition peeling operation. On the research
project of “Replacement of Neanderthals by Modern
Humans: Testing Evolutionary Models of Learning”,
our research reported analysis result of video images
and analysis of illocutionary acts by interview of thesubject. In the past on this project, we have reported
analysis results using interview the subject about
operation process and thinking points with the detailmotion data and the fabrication characteristics of the161should proceed with consideration of the flakes in
the process of acquisition of② it is predicted toadvance (prediction) the transition state to the processof ② . Our discussions would not touch here in thispaper. This is consideration point using experiential
knowledge. We should the future work for analysis
and research. The rehearsal experiment is 1time. Then,
we conducted a full-scale experiment. After lunch,
I conducted a final experiment. LP motion is trying
behavior to get the object have a projectile point. Thisobject are such as a spear, dart, or arrow, or perhaps
used as a knife. If the knapper has an image thatbehavior, the knapper makes an announcement "LP"in the experiment. LF motion is trying behavior to get
the flakes objects. If the knapper has an image thatbehavior, the knapper makes an announcement "LF"
in the experiment. In the rehearsal, the knapper tried
5 times knap for LP / LF. This case, total knap is 28
times during 11 minutes. In the first experiment, theknapper tried 14 times knap for LP / LF. This case,total knap is 52 times during 27 minutes. In the final
experiment, the knapper tried 16 times knap for LP /LF. This case, total knap is 66 times during 55 minutes.
The knapper was carefully shaping the stone. After
that, the knapper hit strongly. Is this same process of
Neanderthal? or Modern humans?

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162

Analysis about getting to LF motion
In this paper, we have processed a detailed analysis
of behavior. Our focusing is the operation at the timeof getting flakes. Figure 2 is the marker position andhummer stone in this experiment. When knapper
gives us declaration for getting flakes, he always
selected most lightweight stone, 115[g]. This is likely
to be a result of determining the controllable easy
than the power impact. Additionally, No.3 marker isrepresenting the movement of the stone. Figure 3 isan example of a photo. Before the motion, knapper
declared the process of getting LF. We would be able
to see the behavior twisting the wrist, and then see
the behavior the pull behind the elbow. Figure 4 is agraph of impulse on the hammer stone at getting FL.
This impulse curve explains the big impact and hittingstone. After this impact, stone speed has changed
slowly. Therefore for getting LF, the knapper takes adecision this impulse value necessary. See Figure 4
and 5, the direction of the impulse is changing rapidly
between about 0.35-0.55 [sec]. This area is a curveafter the stone hammer collides with the stone. The
impulse reaches to target stone by the momentum
conservation law. It shows that the knapper isaccelerating at the same time pulling the elbow andtwist of the wrist. Figure 6 shows the torque applied tothe hammer stone vector. Near the impact point, thetorque vector is a direction different from the directionof travel of the hammer stone. We let discuss this area.
We can consider the vector direction of this graph isdue to a twist of the wrist. In order to obtain a constant
speed of this very short time, twisting of the knapper
operation is to take full advantage of the humanskeleton. This twist makes a small shift of hit points. A
greater speed is controllable easy for the small skeleton.
If the skeleton is strong, the ​​twist operation increases
a shift of hit points. In order to suppress the deviationthat knapper must slow down. If the hammer stoneheavier then, knapper can make the impulse of thesame degree. Skeletal differences make a difference
hummer selection and hitting speed to target stone.
In other words, 3D graph and light weight hummerstone explain that modern small person represents theclever control arm than the strong skeleton people like
Neanderthal.
Discussion
We are considering that Homo sapiens have survived
by twisting motion and the clever control arm. Thisis a necessary element of both individual learningand social learning. To determine the LP and LF
motion is particularly important individual learning.
The individual learning is the most important andnecessary long time learning. This learning method ishighly suitable for modern people life. We assumed toassist the operation of swinging the arm acceleration
to pull the elbow. Neanderthals were 80-90 kg bodyweight, height 160 cm. Their muscles were welldeveloped. They also had strong bones. We speculatethat they are inferior the ability of learning and did
not control the twist of the arm. Therefore, we believethat they were making crushed stone instinctively as
the recurrent Levallois technique. In addition, thelife expectancy of the Neanderthals is about 20 years
old, they were short-lived. The Knowledge accession
to the next generation has been hard. In the future,
we will proceed with the investigation and analysis of
the literature. We will be looking for a material that
represents the difference between the learning ability
between Neanderthal and modern human.

Figure 2. Motion marker position and stone.

Figure 3. Motion pictures of hitting strongly.

Figure 4. Impulse curve getting LF.

Figure 5. 3D acceleration curve No.3 marker.

Figure 6. Direction vector of the rotational
torque around hammer-stone.

Neuroscience Group C02

Figure 1. Two processes of Levallois technique.

163

Brain Activation Related to the Imitative Learning of Bodily
Actions Observed during the Construction of a Mousterian
Stone Tool: A Functional Magnetic Resonance Imaging Study
Naoki Miura1, Kenji Nagai2, Mika Yamazaki3, Yumiko Yoshida4,5,
Hiroki C. Tanabe4,6, Takeru Akazawa7, and Norihiro Sadato4,5

1Department of Information and Communication Engineering, Faculty of Engineering, Tohoku Institute of Technology, Japan
2Department of Historic Heritage, Tohoku University of Art & Design, Japan
3Child Development Research Center, Graduate School of Medical Sciences, National University Corporation University of Fukui, Japan
4Division of Cerebral Integration, Department of Cerebral Research, National Institute for Physiological Sciences, Japan
5Department of Physiological Sciences, School of Life Sciences, The Graduate University for Advanced Studies (Sokendai), Japan
6Graduate School of Environmental Studies, Nagoya University, Japan
7Research Institute, Kochi University of Technology, Japan

Neuroscience Group C02

164

Introduction
An elucidation of cognitive abilities
under pinning social learning behavior areimportant to comprehend the human evolutionary
history, especially for inheritance of technology,
because inheritance of technology is realized bylearning the information about the behavior ofother individuals and its products through several
kinds of social interactions among individuals
such as imitation of the others’ behavior. Previous
studies suggested that the mirror neuron system
including Broca’s area commonly participates inneural basis of tool-use and language, accordingly
it is considered that the neural basis of thosecognitive function have common developmental
process. However, there is no direct evidence
about which component of neural basis of tool-use
played a key role in the process of social learning toacquire novel technology from others. Especially, it
is unknown in whether an acquisition of cognitive
ability for tool-use via the social interaction onprehistorian age was intervened in the learning
ability for linguistic information. In this study,
we focused a neural mechanism to learn a novel
technology such as stone tool-making throughan observation of others’ behavior, and existenceof shared neural mechanism with learning of
linguistic information.
We hypothesize that the stone tool-making canbe learned by observation of the others behavior,and that neural mechanism contains different

neural components compared with the imitative
learning of spoken language. To verify the
hypothesis, we used functional magnetic resonanceimaging. The experimental task was consisted arepetitive observation of the stone tool-making,and that of unknown word-pronunciation.
Materials and Methods
Twenty-four healthy Japanese volunteers
participated in this study (12 males and 12
females, mean age: 25 ± 5, range: 20–36 years).
The experimental data from six subjects were
excluded because of excessive head movement
or an insufficient number of responses. Thus, weanalyzed data from 18 participants (nine males andnine females). All participants provided written
informed consent to an experimental protocol
approved by the Ethical Committee of the
National Institute for Physiological Sciences. The
authors also declare that all experiments on humansubjects were conducted in accordance with the
Declaration of Helsinki.
The fMRI experiment consisted of four runsof actual measurement and one practical run. A
rapid event-related design was used for the fMRI
experiment. During fMRI session, all participants
observed 15 moving pictures and one still pictureof stone tool-making and 15 moving picturesand one still picture of the pronunciation of an
Uzbek word. Each picture showed a kind of bodily
action used to make a Mousterian stone tool or toResults and Discussion
We found task-specific cortical activation of
the parietofrontal network during the observationof stone tool-making. A recent meta-analysisof action-observation and imitation tasks also
reported similar patterns in cortical networks(Caspers et al., 2010). The results suggest thatthe cognitive mechanism to manipulate theself-motor representations contributes in theaction observation during the imitative learning.
By contrast, the bilateral premotor area andsuperior temporal gyrus were activated during
the observation of word-pronunciation. Previous
neuroimaging studies of phonological processing
reported that the inferior part of the dorsal
premotor area plays an important role in thearticulatory process (Brown et al., 2009), and theposterior part of the superior temporal gyrus
is important for audiovisual speech perception(Murase et al., 2008), thus supporting ourconclusion that the activation we observed in thebilateral superior temporal gyrus was induced byperceptions of the pronunciation of the Uzbek
word.
The learning-related activation change duringobserving the stone tool-making was shownin the right cerebellum (Figure 1(a)) and thatof word-pronunciation was shown in the left
superior temporal gyrus (Figure 1(b)). The result
suggested that the progress of imitation learning
was represented on the specific cortical region,and the represented region was dependent on theinformation which the subject focused to learn. A
previous study reported that the changes of activity
in cerebellum reflects a progress of internal model
formulation to learn a motor control to manipulate
a novel tool (Imamizu et al., 2000), whereas therepetition-related decreases in the left superior
temporal gyrus reflected the effect of learning on
the neural processing of the perception of auditory
stimuli (Rauchecker et al., 2008). By contrast, thecommon activated regions of both observation
tasks are not influenced by progress of imitative
learning.
In conclusion, the progress of imitative learningon stone tool-making and word–pronunciation
were represented activity of different neuralmechanism instead of common neural mechanism
between them. Therefore, the result indicatethat the progress of imitation learning for thestone tool-making and word-pronunciation

Neuroscience Group C02

pronounce one Uzbek word. Each moving picture
was presented twice in each run. The practical run
presented moving pictures that differed from thoseused in the actual run. The pictures were separated
by resting intervals of approximately 4 sec, during
which time a white fixation cross was presented.
Participants were instructed to observe thepictures and to memorize the content of the bodily
action or word pronunciation. To ensure that theparticipants were conscious of the task, an actual
imitation task was conducted immediately after
the fMRI measurement. To confirm the arousal
state of the participants, the color of the fixation
cross occasionally changed to yellow, and theparticipants were instructed to press a button when
they noticed the change. Figure 1 illustrates atimeline of an fMRI run.
Data preprocessing and statistical analysesof fMRI data were performed using statisticalparametric mapping (SPM8, Wellcome TrustC e n t e r f o r Ne u ro i m a g i n g, L on d on , U K ) .
The fMRI data were analyzed using a twostage approach. D uring the first stage, thehemodynamic responses produced under thedifferent experimental conditions were assessedat each voxel on an intra-subject basis using ageneral linear model. The second stage of analysis
was performed on an inter-subject basis using a
two-way repeated-measures factorial design. One
factor was the type of picture observed, and theother factor was the number of times the samemoving picture was presented repeatedly. To
identify the regions showing learning effect foreach task, contrast images representing a repetition
suppression effect of task-related activation werecreated and estimated. The statistical threshold
was set at p < 0.05 (corrected for family-wise error(FWE) by voxel level).

165

were represented on the specific cortical region
respectively, and the represented region wasdependent on the information which the subject
focused to learn.
References
Caspers, S., Zilles, K., Laird, A.R., Eickhoff, S.B. 2010
ALE meta-analysis of action observation andimitation in the human brain. Neurimage 50: 1148-67.
Brown, S., Laird, A.R., Pfordresher, P.Q., Thelen, S.M.,
Turkeltaub, P., Liotti, M. 2009 The somatotopy ofspeech: Phonation and articulation in the humanmotor cortex. Brain and Cognition 70: 31-41.
Imamizu, H., Miyauchi, S., Tamada, T., Sasaki, Y., Takino,
R., Puetz, B., Yoshioka, T., Kawato, M. 2000 Human

cerebellar activity reflecting an acquired internal
model of a novel tool. Nature 403:192-195.
Murase, M., Saito, D.N., Kochiyama, T., Tanabe,
H.C., Tanaka, S., Harada, T., Aramaki, Y., Honda,
M., Sadato, N. 2008 Cross-modal integration
during vowel identification in audiovisual speech:
A functional magnetic resonance imaging study.
Neuroscience letters 434: 71-76.
Rauschecker, A.M., Pringle, A., Watkins, K.E. 2008
Changes in neural activity associated with learning
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Sato, W., Kochiyama, T., Yoshikawa, S., Naito, E.,
Matsumura, M. 2004 Enhanced neural activity inresponse to dynamic facial expressions of emotion: an
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Figure 1. Cortical region showing repetition-suppression effect in the (a) right cerebellar posterior lobule with taskrelated activation during observation of stone-tool making, and (b) in the left superior temporal gyrus with task-related
activation during observation of word pronunciation.

Neuroscience Group C02

166

Abe-Ouchi, A.

6, 66, 143, 164

Amano, H.

66, 151, 159

Ando, J.

97

Aoki, K.

56, 58

Bar-Yosef, O.

Author Index

42, 137

Akazawa, T.

24

Barkai, R.

35

Bortolini, M. C.

135

Boyette, A. H.

99

Bruner, E.

68

Chan, W.-L.

137

Creanza, N.

27

Diab, M. C.

139

Feldman, M. W.

27

Fogarty, L.

27

Gopher, A.

35

Hagino, I.

101, 119

Haizuka, Y.

72

Hatano, A.

61

Hayashi, K.

103

Hewlett, Bonnie

105

Hewlett, Barry S.

55

Higurashi, Y.

86

Hirai, N.

72

Hishida, H.

153

Horiuchi, S.

53, 121

Hoshino, Y.

161

Hünemeier, T.

135

Imamura, K.

108

Inaba, M.

61

Ishida, H.

66

Ishii, R.

103

Kadowaki, S.

38

Kamei, N.

110

Kato, H.

40

Kawahata, H.

42, 145

Kawamichi, H.

79

Kawasaki, K.

123

Oguchi, T.

42, 141, 149

Kikuchi, T.

66

Ohtake, Y.

153

Kimura, R.

125

Omori, T.

147

Kitada, R.

79

Omura, K.

114

Kobayashi, Y.

72, 127

Onoda, R.

61

Kochiyama, T.

74

Oota, S.

153

Kondo, O.

66, 70, 151, 153, 157, 159

Ponce de León, M.

30

Kondo, Y.

88, 141, 149

Paixão-Côrtes, V. R.

135

112

Koyama, T.
Kubo, D.

70, 155

Sadato, N.

79, 164

Salzano, F. M.

135
38

Kubota, S.

53

Sano, K.

Kuhn, S. L.

44

Sasaki, A.

79

Lehmann, L.

58, 60

Shimada, M.

133

Marukawa, Y.

143

Simunovic, D.

61

Matsui, T.

72

Song, W.

149

Matsumura, G.

72

Stout, D.

78

Matsunaga, M.

79

Suzuki, H.

66, 151, 153, 157

63

Takada, A.

116

Mesoudi, A.
Michikawa, T.

66, 153, 157

Takahashi, H.

79

Mithen, S. J.

28

Takahashi, K.

137

Miura, N.

161, 164

Takahashi, N.

61

Mizuno, K.

82

Takakura, J.

48, 94Mori, H.

143

Tanabe, H. C.

74, 76, 155, 164

Moriguchi, M.

157

Terashima, H.

45, 103

Morita, Y.

66, 151, 159

Viscardi, L. H.

135

Nagai, K.

161, 164

Wakano, J. Y.

58

151

Yamagami, E.

117

Naganuma, M.

40

Yamauchi, T.

101, 119

Nakahashi, W.

130

Yamazaki, M.

164

Nakamura, Yoshifumi

143

Yokoyama, Y.

42, 145

Nakamura, Yuuki

103

Yoneda, M.

42, 139, 147

Nakata, H.

90

Yoshida, Y.

79, 164

Nagano, H.

Nakatsukasa, M.

66, 151, 159

Yoshihara, K.

79

Nishiaki, Y.

6, 50, 92, 103Zilhão, J.

34

Zollikofer, C.

30

O'ishi, R.

137

Obrochta, S.

145

Ogihara, N.

66, 70, 72, 74, 151, 153, 157, 159

RNMH2012
The First International Conference November 18-24, 2012, Tokyo
Edited by Takeru Akazawa & Yoshihiro Nishiaki

RNMH PROJECT SERIES No. 002

RNMH2012
The First International Conference

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

November 18-24, 2012, Tokyo
Edited by

Takeru Akazawa &Yoshihiro Nishiaki