From Biped to Strider The Emergence ofModem Human Walking, Running, andResource Transport From Biped to Strider 0/ The Emergence Modern Human Walking, Running, and Resource Transport Edited by D. Jeffrey Meldrum ldaho State University Pocatello, Idaho and Charles E. Hilton Western Michigan University Kalamazoo, Michigan Springer-Science+Business Media, LLC Library of Congress Cataloging-in-Publication Data From biped to strider : the emergence of modem human walking, running, and resource transport I edited by D. Jeffrey Meldrum and Charles E. Hilton. p. cm. Based on a symposium which was held as a session of the 69th annual meeting of the AAPA held in San Antonio, Texas, in 2000. Includes bibliographical references and index. ISBN 978-0-306-48000-3 ISBN 978-1-4419-8965-9 (eBook) DOI 10.1007/978-1-4419-8965-9 1. Human locomotion-Congresses. I. Meldrum, D. Jeffrey, 1958- II. Hilton, Charles E. III. American Association of Physical Anthropologists. Meeting (69th: 2000 : San Antonio, Tex.) QP303.F685 2004 612.7'6--dc22 2003060287 ISBN 978-0-306-48000-3 © 2004 Springer Science+Business Media New York Originally published by Kluwer AcademicIPlenum Publishers, New York in 2004 Softcover reprint ofthe hardcover 1st edition 2004 10 9 8 7 6 5 4 3 2 1 A C.LP. record for this book is available from the Library of Congress. All rights reserved No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Permissions for books published in Europe: [email protected] Permissions for books published in the United States of America: [email protected] Participantsinthesymposium,FromBipedtoStrider:TheEmergenceofModernHumanWalking,heldatthe 69thannualmeetingoftheAAPAheldinSanAntonio,Texas,April 15,2000.Frontrow(Itor):Roshna Wunderlich,LauraGruss,MarshaOgilvie,KarenRosenberg,DavidBegun,JeffMeldrum.Backrow(Itor): DennisBramble,ChrisRuff,Chuck Hilton,DavidCarrier [notpictured:RonClarke,YvetteDeloison, PeterSchmid,DanSchmitt,PatriciaKramer]. Contributors DavidR. Begun, DepartmentofAnthropology, University ofToronto,Toronto,ON M5S 3G3,Canada Gilles Berillon,CNRS, Paris,France DavidCarrier,DepartmentofBiology, University ofUtah, Salt LakeCity,UT84112 YvetteDeloison, CNRS, Paris,France RussellD.Greaves, Department ofAnthropology, University ofTexas,SanAntonio, TX 78249 Laura Tobias Gruss, Department of Biological Anthropology & Anatomy, Duke University,Durham, NC27708 Cbarles E. Hilton, Department of Anthropology, Western Michigan University, Kalamazoo,MI49008 Patricia Ann Kramer, Departments of Anthropology and Orthopaedics, University of Washington,Seattle,WA98104-2499 Henry M McHenry, Department of Anthropology, University of California, Davis, CA 95616 D.JeffreyMeldrum,DepartmentofBiologicalSciences,IdahoStateUniversity,Pocatello, ID83209-8007 MarshaD.Ogilvie,DepartmentofAnthropology,UniversityofNewMexico,Albuquerque, NM87131 Peter Schmid, Anthropology Institute, University of Zurich-Irchel, Zurich CH8057, Switzerland Daniel Schmitt, Department of Biological Anthropology & Anatomy, Duke University, Durham, NC27708 vii Preface Theinspirationforthisvolumeofcontributedpapersstemmedfromconversationsbetween theeditorsinfrontofChuckHilton'sposteronthedeterminants ofhominidwalkingspeed, presented at thel998 meetings of the American Association of Physical Anthropologists (AAPA).Earlieratthosemeetings,JeffMeldrum (withRoshna Wunderlich)hadpresented analternate interpretationoftheLaetolifootprints basedonevidenceofmidfoot flexibility. As the discussion ensued we found convergence on a number of ideas about the nature of the evolution of modem human walking. From the continuation of that dialogue grew the proposal for a symposium which we called FromBiped to Strider: the Emergence of ModemHuman Walking.Thesymposiumwasheldasasessionofthe69thannualmeeting oftheAAPA,heldinSanAntonio, Texasin2000.Itseemed tousthatthestudyofhuman bipedalismhadbecomeovershadowedbytheoftenpolarizeddebatesoverwhetheraustralo pithecines were wholly terrestrial in habit,or retained a significantdegree of arboreality. Furthermore, there remained considerable difference of opinion as to whether bipedalism aspracticed by australopithecines waseither equivalent to modem human walking,inter mediate innature,orunique.Itseemedtousthatpreoccupationwiththese issues hadlead to adegree ofdisregard concerning the fact that modem human locomotion is more than merelyhabitually walkingontwolegs.Ithasbecomequiteevidentthatofallprimates,and of the majority of mammals,modem humans are notably adapted for endurance walking andrunning.Wefeltthatwiththeadventofthenewmillennium itwastimetoshiftattention tothepatternandtimingoftheemergence ofmodem human walking,characterizedbythe development of the longitudinal arch, midfoot stability, and plantar fascia modifications that accompanied the human striding stiff-legged gait. When and in what sequence did these morphological traits appear? What were the changes in the bio-behavioral complex ofhominin locomotorevolution? What weretheimplications fortheenhancementandthe expansion ofhominin mobility? We invited participants that would represent both paleontological and neotological perspectives on the functional morphology and behavioral ecology of hominin locomotor adaptations.Thesecontributionsspanfromtheoriginsofbipedalismtolocomotorcontrasts among contemporary human populations. We purposely recruited participants largely from theupcoming generationofanthropologistsinorder toinfuse fresh perspectives and insights into the dialogue, while building upon the pioneering researches and discussions of senior colleagues. Their contributions before, during, and after the symposium were thought-provoking, Thisvolumereports theproceedings ofthatsymposium.Thesubtitlewasexpandedto properlyrecognize thesignificantrolesofrunning andresource transportinmodem human locomotor adaptations. Regretably, not all the participants were able to contribute to the ix x Preface volume, but we also picked up an additional paper along the way from Gilles Berrilion, and Henry McHenry graciously agreed to contribute a discussion paper. The papers in this volume were reviewed by at least three reviewers and the overall quality benefited greatly from theexpertise andprofessionalism displayedbyeach ofthem.Wealso wish to acknowledgethe long-sufferingandassistanceofourpublisher, Teresa Krauss. In the time between the symposium and the completion of this volume, new fossil discoveries continuetoaddpiecestothepuzzleofhumanlocomotorevolution. Atthesame time,theapplicationofnewinvestigatorytechniquesandbroaderinterdisciplinaryperspec tivesprovide refinedinterpretationsofthepaleontologicalandneontologicalevidence. Itis ourhopethatthisvolume willspawnnovelperspectivesandpenetratingquestionsdirected at these issues,and that it will contribute to the ongoing dialogue between those who are interestedinthehominin transitionfrom biped tostrider. JeffMeldrum Chuck Hilton Contents Chapter1 Striders,Runners, and Transporters..................................................... 1 Charles E.Hilton and D.JeffreyMeldrum Chapter2 Knuckle-Walkingand the OriginofHumanBipedalism......................... 9 DavidR.Begun Chapter3 ANewHypothesis on the OriginofHominoid Locomotion...................... 35 Yvette Deloison Chapter4 FunctionalInterpretationofthe Laetoli Footprints................................. 49 PeterSchmid ChapterS FossilizedHawaiianFootprints Compared with Laetoli Hominid Footprints............................................................................. 63 D. JeffreyMeldrum Chapter6 In WhatMannerDidThey Walk onTwoLegs? AnArchitectural Perspectivefor the FunctionalDiagnosticsofthe EarlyHominidFoot..... 85 Gilles Berillon Chapter7 The BehavioralEcologyofLocomotion 101 PatriciaAnn Kramer Chapter8 Bipedalism inHomo ergaster: AnExperimentalStudy ofthe Effects ofTibial Proportionson LocomotorBiomechanics 117 Laura TobiasGruss and DanielSchmitt xi xii Contents Chapter9 The Running-Fighting Dichotomy and the Evolution ofAggression in Hominids 135 DavidCarrier Chapter10 Age,Sex,and Resource Transportin Venezuelan Foragers 163 Charles E.Hilton andRussellD. Greaves Chapter11 Mobilityand the LocomotorSkeletonat the Foraging to FarmingTransition 183 Marsha D. Ogilvie Chapter12 Uplifted Head,FreeHands, and the Evolution ofHuman Walking 203 Henry M. McHenry Index 211 From Biped to Strider The Emergence ofModern Human Walking, Running, andResource Transport