s t o n e a g e i n s t i t u t e p u b l i c a t i o n s e r i e s Series Editors Kathy Schick and Nicholas Toth Stone Age Institute Gosport, Indiana and Indiana University, Bloomington, Indiana Number 1. THE OLDOWAN: Case Studies into the Earliest Stone Age Nicholas Toth and Kathy Schick, editors Number 2. BREATHING LIFE INTO FOSSILS: Taphonomic Studies in Honor of C.K. (Bob) Brain Travis Rayne Pickering, Kathy Schick, and Nicholas Toth, editors Number 3. THE CUTTING EDGE: New Approaches to the Archaeology of Human Origins Kathy Schick, and Nicholas Toth, editors Number 4. THE HUMAN BRAIN EVOLVING: Paleoneurological Studies in Honor of Ralph L. Holloway Douglas Broadfield, Michael Yuan, Kathy Schick and Nicholas Toth, editors ii 3 The Human Brain Evolving: Papers in Honor of Ralph L. Holloway T S A I P he Tone ge nSTITuTe reSS P S ublIcATIon erIeS c S e ommenT by erIeS dITorS The Stone Age Institute Press has been established Institute include organizing workshops - convening top to publish critical research into the archaeology of hu- scientists to discuss and work together on special topics man origins. The Stone Age Institute is a federally-ap- - and organizing and sponsoring conferences, lectures, proved non-profit organization whose mission is investi- and internet websites to disseminate information to a gating and understanding the origins and development of broader, general public. human technology and culture throughout the course of An equally critical component of our operations is human evolution, and to disseminate information to the the publication of research results in journal articles and general public regarding science education, in particu- books. The Human Brain Evolving: Paleoneurological lar about our evolutionary origins. The ultimate goal of Studies in Honor of Ralph L. Holloway is the fourth vol- these undertakings is to provide a better understanding ume in a new publication series initiated by the Stone of the human species and our place in nature. Age Institute Press. This volume emanates from a Stone An unbroken line of technology and culture extends Age Institute conference in honor of the lifework of from our present day back in time at least two-and-a- Ralph Holloway and brain evolution. This meeting con- half million years. The earliest stone tool-makers were vened researchers at Indiana University and the Stone upright-walking, small-brained ape-men, and from these Age Institute for two days in April of 2007 to present a primordial origins, the human lineage embarked upon a wide variety of research pertaining to the evolution of pathway that is extraordinary and unique in the history the human brain. The Stone Age Institute Press is de- of life. This evolutionary pathway has shaped our bod- voted to the publication of seminal research in the form ies, our brains, and our way of life, and has had an ever- of high-quality, data-rich volumes, including informa- accelerating impact on the earth and its other organisms. tion and interpretations from original research as well as Well over 99 percent of the time span of this techno- critical syntheses of research results. logical evolution occurred in the Stone Age, so it is no For more information on the Stone Age Institute, in- exaggeration to say that modern humans have evolved as cluding its research programs, lectures, conferences, and Stone Age creatures. publications, we invite you to visit our web site at www. The Stone Age Institute’s goal is to provide a ha- stoneageinstitute.org. To explore our most recent sci- ven for this research, providing support for scholars ence education project on evolution and “Big History,” from around the world conducting cutting-edge human visit the web site www.fromthebigbang.com. origins research. Our research team and associates in- clude professional research scientists in long-term posi- tions, postdoctoral research fellows, and visiting schol- ars. Fieldwork, collecting primary data from important Kathy Schick and Nicholas Toth archaeological sites, as well as laboratory studies and Co-Directors, Stone Age Institute experimental archaeological research are all critical Series Editors, Stone Age Institute Press Publication components of our operation. Other vital functions of the Series S T O N E A G E I N S T I T U T E P U B L I C AT I O N S E R I E S N U M B E R 4 Series Editors Kathy Schick and Nicholas Toth THE HUMAN BRAIN EVOLVING: Paleoneurological Studies in Honor of Ralph L. Holloway Editors Douglas Broadfield Florida Atlantic University Michael Yuan Columbia University Kathy Schick Stone Age Institute & Indiana University Nicholas Toth Stone Age Institute & Indiana University Stone Age Institute Press · www.stoneageinstitute.org 1392 W. Dittemore Road · Gosport, IN 47433 FRONT COVER CAPTIONS Center: Portrait of Ralph L. Holloway. Upper left: A modern human brain. Upper right: Ralph measuring landmarks on an endocast ca. 1976. Lower right: Homo habilis cranium KNM-ER-1813 from Koobi Fora, Kenya (photo by Holloway). Lower left: Ralph with an endocast of the Flores “hobbit” cranium. Published by the Stone Age Institute. ISBN-10: 0-9792276-3-1 ISBN-13: 978-0-9792276-3-9 Copyright © 2010, Stone Age Institute Press. All right reserved under International and Pan-American Copyright Conventions. No part of this book may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, without permission in writing from the publisher. d edIcATIon To Michael Sheng-Tien Yuan Michael Sheng-Tien Yuan, D.D.S., M.A., M.S., sertation work was on dental development, Michael is Ph.D. was born in Taiwan (ROC) on November 17, most remembered in anthropology for his contributions 1959. In an earlier life before he chose orthodontics and to our understanding of human brain evolution. After anthropology as his careers, Michael was an internation- completing his Ph.D. in 2000, Michael moved back to ally recognized actor, earning acclaim for his role in the Columbia’s College of Dental Medicine as an Assistant Taiwanese motion picture Jade Love (1984), which re- Professor of Clinical Dental Medicine. As with every- ceived a number of Golden Horse Awards and was fea- thing he did Michael threw himself entirely into his new tured at film festivals around the globe. In 1988 after profession, anatomist. He earned a third dental degree completing his dental degree at National Taiwan Uni- in 2003. versity and two years of compulsory military service, In his short tenure at Columbia, Michael became Michael moved to New York to work on his Master’s in one of the most beloved instructors in not only the Col- orthodontics from Columbia University. After complet- lege of Medical Dentistry, but also Columbia’s College ing his orthodontics degree Michael entered the Ph.D. of Physicians and Surgeons, earning three Teacher of the program in Anthropology at Columbia. Shortly after Year awards. He was promoted to Associate Professor entering the anthropology program Michael had his last in 2008. After Michael became ill he continued to teach brush with the acting bug when he was offered the lead from his hospital bed, sending the medical and dental in Ang Lee’s first American film, The Wedding Banquet students details of the various procedures he underwent, (1993). Michael turned down the offer, choosing instead and relating them to the current anatomical region the to focus on his new career. students were learning. True to the way he lived, the Michael was Ralph Holloway’s student from the dedication on his last teaching award summarizes our moment he entered the anthropology program. He feelings for Michael: …in appreciation for his “wisdom, worked closely with Ralph and Doug Broadfield on a gentleness, and ability to … accept life’s challenges and number of endocast projects, bringing his ample artistic use them to grow.” skills to the science of paleoneurology. Though his dis- viii 3 The Human Brain Evolving: Papers in Honor of Ralph L. Holloway Contributors 4 ix Contributors 4 ix c onTrIbuTorS John M. Allman ([email protected]) Division of Biology, California Institute of Technology, Pasadena, California, USA Nicole Barger ([email protected]) Department of Anthropology, University of California at San Diego, USA Doug Broadfield ([email protected]) Department of Anthropology, Florida Atlantic University, Boca Raton, Florida, USA Emiliano Bruner ([email protected]) Centro Nacional de Investigación sobre la Evolución Humana, Burgos, Spain Daniel P. Buxhoeveden ([email protected]) Department of Psychology, University of South Carolina, Columbia, South Carolina, USA Joseph M. Erwin ([email protected])Foundation for Comparative and Conservation Biology, Needmore, PA Dominique Grimaud-Herve ([email protected]) Département de Préhistoire, Muséum national d'Histoire naturelle, Paris, France John W.K. Harris ([email protected]) (PhD, U California-Berkeley, 1978; Prof, SAS) Center for Human Evolutionary Studies, Department of Anthropology, Rutgers University, New Brunswick, New Jersey, USA Patrick R. Hof ([email protected]) Department of Neuroscience, Mount Sinai School of Medicine, New York, NY Ralph L. Holloway ([email protected]) Department of Anthropology, Columbia University, New York, New York, USA Karin Isler ([email protected]) Anthropological Institute & Museum, University of Zürich- Irchel, Zürich Jason A. Kaufman ([email protected]) Division of Biology, California Institute of Technology, Pasadena, California, USA James E. King ([email protected]) Emeritus, Department of Psychology, University of Arizona, Tucson, Arizona, USA David Lordkipanidze Georgian National Museum, Departement of Geology and Paleontology, Academy of Sciences of Georgia, Tbilissi, Georgia Alan Mann ([email protected]), Department of Anthropology, Princeton University, Princeton, New Jersey, USA Robert D. Martin ([email protected]) Department of Anthropology, The Field Museum, Chicago, Illinois, USA Janet Monge ([email protected]), Department of Anthropology, University of Pennsylvania, Philadelphia, Pennsylvania, USA Francine “Penny” Patterson The Gorilla Foundation, Woodside, California, USA Mary Ann Raghanti ([email protected]) Department of Anthropology and School of Biomedical Sciences, Kent State University, Kent, OH x 3 The Human Brain Evolving: Papers in Honor of Ralph L. Holloway Contents 4 xi James K. Rilling ([email protected]) Department of Anthropology and Center for Behavioral Neuroscience, Emory University, Department of Psychiatry and Behavioral Sciences, Emory University School of Medicine and Division of Psychobiology, Yerkes National Primate Research Center, Atlanta, Georgia, USA Duane M. Rumbaugh ([email protected]) Emeritus, Language Research Center and Department of Psychology, Georgia State University, Atlanta, Georgia, USA E. Sue Savage-Rumbaugh ([email protected]) Great Ape Trust, Des Moines, Iowa, USA Natalie Schenker ([email protected] ) Department of Anthropology, University of California at San Diego and Department of Anthropology, The George Washington University, Washington, D.C., USA Kathy Schick ([email protected]) Stone Age Institute and Indiana University, Bloomington, Indiana, USA P. Tom Schoenemann ([email protected]) Anthropology Department, Indiana University, Bloomington, Indiana and Stone Age Institute, USA Katerina Semendeferi ([email protected] ) Department of Anthropology and Graduate Program in Neurosciences, University of California at San Diego, USA Chet C. Sherwood ([email protected]) Department of Anthropology, The George Washington University, Washington, D.C. Francys Subiaul ([email protected]) The George Washington University, Mind, Brain & Evolution Center, Department of Speech & Hearing Science, Washington, D.C., USA Jared Taglialatela ([email protected], [email protected]) Department of Biology, Kennesaw State University and Yerkes National Primate Research Center, Emory University, Atlanta, Georgia, USA Nicholas Toth ([email protected]) Stone Age Institute and Indiana University, Bloomington, Indiana, USA J. Michael Tyszka ([email protected]) Brain Imaging Center, California Institute of Technology, Pasadena, California, USA Eric J. Vallender ([email protected]) New England Primate Research Center, Harvard Medical School, Southborough, Massachusetts, USA. Anne H. Weaver, Ph.D. ([email protected]) Anthropology Department, University of New Mexico, Albuquerque, New Mexico, USA