Thumbnail · 2015. 10. 27. · The Blackwell Companions to Anthropology offers a series of comprehensive syntheses of the traditional subdisciplines, primary subjects, and geographic

A Companion to Dental Anthropology

The Blackwell Companions to Anthropology offers a series of comprehensive syntheses of the traditional subdisciplines primary subjects and geographic areas of inquiry for the field Taken together the series represents both a contemporary survey of anthropology and a cutting edge guide to the emerging research and intellectual trends in the field as a whole

1 A Companion to Linguistic Anthropology edited by Alessandro Duranti2 A Companion to the Anthropology of Politics edited by David Nugent and Joan

Vincent3 A Companion to the Anthropology of American Indians edited by Thomas Biolsi4 A Companion to Psychological Anthropology edited by Conerly Casey and Robert

B Edgerton5 A Companion to the Anthropology of Japan edited by Jennifer Robertson6 A Companion to Latin American Anthropology edited by Deborah Poole7 A Companion to Biological Anthropology edited by Clark Larsen (hardback only)8 A Companion to the Anthropology of India edited by Isabelle Clark‐Decegraves9 A Companion to Medical Anthropology edited by Merrill Singer and Pamela

I Erickson10 A Companion to Cognitive Anthropology edited by David B Kronenfeld Giovanni

Bennardo Victor de Munck and Michael D Fischer11 A Companion to Cultural Resource Management edited by Thomas King12 A Companion to the Anthropology of Education edited by Bradley AU Levinson

and Mica Pollack13 A Companion to the Anthropology of the Body and Embodiment edited by Frances

E Mascia‐Lees14 A Companion to Paleopathology edited by Anne L Grauer15 A Companion to Folklore edited by Regina F Bendix and Galit Hasan‐Rokem16 A Companion to Forensic Anthropology edited by Dennis Dirkmaat17 A Companion to the Anthropology of Europe edited by Ullrich Kockel Maacuteireacutead

Nic Craith and Jonas Frykman18 A Companion to Border Studies edited by Thomas M Wilson and Hastings Donnan19 A Companion to Rock Art edited by Jo McDonald and Peter Veth20 A Companion to Moral Anthropology edited by Didier Fassin21 A Companion to Gender Prehistory edited by Diane Bolger22 A Companion to Organizational Anthropology edited by D Douglas Caulkins

and Ann T Jordan23 A Companion to Paleoanthropology edited by David R Begun24 A Companion to Chinese Archaeology edited by Anne P Underhill25 A Companion to the Anthropology of Religion edited by Janice Boddy and Michael

Lambek26 A Companion to Urban Anthropology edited by Donald M Nonini27 A Companion to the Anthropology of the Middle East edited by Soraya Altorki28 A Companion to Heritage Studies edited by William Logan Maacuteireacutead Nic Craith

and Ullrich Kockel29 A Companion to Dental Anthropology edited by Joel D Irish and G Richard Scott

A Companion to Dental AnthropologyEdited byJoel D Irish and G Richard Scott

This edition first published 2016copy 2016 John Wiley amp Sons Inc

Registered OfficeJohn Wiley amp Sons Ltd The Atrium Southern Gate Chichester West Sussex PO19 8SQ UK

Editorial Offices350 Main Street Malden MA 02148‐5020 USA9600 Garsington Road Oxford OX4 2DQ UKThe Atrium Southern Gate Chichester West Sussex PO19 8SQ UK

For details of our global editorial offices for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at wwwwileycomwiley‐blackwell

The right of Joel D Irish and G Richard Scott to be identified as the authors of the editorial material in this work has been asserted in accordance with the UK Copyright Designs and Patents Act 1988

All rights reserved No part of this publication may be reproduced stored in a retrieval system or transmitted in any form or by any means electronic mechanical photocopying recording or otherwise except as permitted by the UK Copyright Designs and Patents Act 1988 without the prior permission of the publisher

Wiley also publishes its books in a variety of electronic formats Some content that appears in print may not be available in electronic books

Designations used by companies to distinguish their products are often claimed as trademarks All brand names and product names used in this book are trade names service marks trademarks or registered trademarks of their respective owners The publisher is not associated with any product or vendor mentioned in this book

Limit of LiabilityDisclaimer of Warranty While the publisher and authors have used their best efforts in preparing this book they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom If professional advice or other expert assistance is required the services of a competent professional should be sought

Library of Congress Cataloging‐in‐Publication Data

A companion to dental anthropology edited by Joel D Irish and G Richard Scott pages cm ndash (Wiley Blackwell companions to anthropology 29) Includes bibliographical references and index ISBN 978-1-118-84543-1 (cloth)1 Dental anthropology I Irish Joel D editor II Scott George Richard editor GN209C66 2015 5999prime43ndashdc23 2015022952

A catalogue record for this book is available from the British Library

Cover image From top to bottom Christy G Turner II Used with permission of the estate of Christy G Turner II Two Neolithic male crania from Gebel Ramlah cemetery Western Desert Egypt Photo by Irish February 2003 Swazi skull with tooth loss tooth tilting alveolar resorption and super-eruption Cat A2306 Dart Collection (University of the Witwatersrand South Africa) Joel D Irish 3-8-2011

Set in 1012pt Galliard by SPi Global Pondicherry India

1 2016

Contents

Notes on Contributors viii

Foreword xv

Acknowledgments xviii

Part I Context 1

1 Introduction to Dental Anthropology 3Joel D Irish and G Richard Scott

2 A Brief History of Dental Anthropology 7G Richard Scott

Part II Dental Evolution 19

3 Origins and Functions of Teeth From ldquoToothedrdquo Worms to Mammals 21Peter S Ungar

4 The Teeth of Prosimians Monkeys and Apes 37Frank P Cuozzo

5 The Hominins 1 Australopithecines and Their Ancestors 52Lucas K Delezene

6 The Hominins 2 The Genus Homo 67Maria Martinoacuten‐Torres and Joseacute Mariacutea Bermuacutedez de Castro

vi contents

Part III The Human Dentition 85

7 Terms and Terminology Used in Dental Anthropology 87Joel D Irish

8 Anatomy of Individual Teeth and Tooth Classes 94Loren R Lease

9 The Masticatory System and Its Function 108Peter W Lucas

Part IV Dental Growth and Development 121

10 An Overview of Dental Genetics 123Toby Hughes Grant Townsend and Michelle Bockmann

11 Odontogenesis 142Edward F Harris

12 Tooth Eruption and Timing 159Helen M Liversidge

13 Tooth Classes Field Concepts and Symmetry 172Grant Townsend Alan Brook Robin Yong and Toby Hughes

Part V Dental Histology from the Inside Out 189

14 The Pulp Cavity and Its Contents 191Scott S Legge and Anna M Hardin

15 Dentine and Cementum Structure and Properties 204Nancy Tang Adeline Le Cabec and Daniel Antoine

16 Enamel Structure and Properties 223Daniel Antoine and Simon Hillson

Part VI Dental Morphometric Variation in Populations 245

17 Identifying and Recording Key Morphological (Nonmetric) Crown and Root Traits 247G Richard Scott Christopher Maier and Kelly Heim

18 Assessing Dental Nonmetric Variation among Populations 265Joel D Irish

19 Measurement of Tooth Size (Odontometrics) 287Brian E Hemphill

20 Assessing Odontometric Variation among Populations 311Brian E Hemphill

Part VII Dental Morphometric Variation in Individuals 337

21 Forensic Odontology 339Heather JH Edgar and Anna LM Rautman

contents vii

22 Estimating Age Sex and Individual ID from Teeth 362Christopher W Schmidt

23 Indicators of Idiosyncratic Behavior in the Dentition 377Christopher M Stojanowski Kent M Johnson Kathleen S Paul and Charisse L Carver

24 Dentition Behavior and Diet Determination 396Kristin L Krueger

Part VIII Dental Health and Disease 413

25 Crown Wear Identification and Categorization 415Scott E Burnett

26 Caries The Ancient Scourge 433Daniel H Temple

27 Dental Stress Indicators from Micro‐ to Macroscopic 450Debbie Guatelli‐Steinberg

28 A Host of Other Dental Diseases and Disorders 465Greg C Nelson

Part IX The Future of Dental Anthropology 485

29 New Directions in Dental Development Research 487John P Hunter and Debbie Guatelli‐Steinberg

30 Chemical and Isotopic Analyses of Dental Tissues 499Louise T Humphrey

31 Non‐Invasive Imaging Techniques 514Joseacute Braga

Index 528

Notes on Contributors

Daniel Antoine is the British Museumrsquos Curator of Physical Anthropology with responsibility for the museumrsquos human remains He completed his PhD at University College London in 2001 and has published widely on bioarchaeology and dental anthropology including Ancient Lives New Discoveries Eight Mummies Eight Stories (2014) with John Taylor and Regarding the Dead Human Remains in the British Museum (2014) with Alexandra Fletcher and JD Hill

Joseacute‐Mariacutea Bermuacutedez de Castro PhD in Biology and Research Professor of the Consejo Superior de Investigaciones Cientiacuteficas (CSIC Spain) is currently affiliated with the National Research Center on Human Evolution (Burgos Spain) From 1991 he has been co‐director of the Sierra de Atapuerca project His main scientific contributions deal with the study of the hominin fossils recovered from the Sierra de Atapuerca sites including the naming in Science of a new Homo species Homo antecessor

Michelle Bockmann is a member of the Craniofacial Biology Research Group in the School of Dentistry at the University of Adelaide Australia Her main research interests involve studies of the human dentition aimed at clarifying the roles of genetic epigenetic and environmental factors on dental development and morphology together with trans-lation of research findings to the general community

Joseacute Braga is Professor of Evolutionary Anthropology and Human Evolution in the Research Department of Biological Anthropology of the University of Toulouse France He conducts fieldwork in Kromdraai (South Africa) where he recovers fossil remains of human ancestors and their relatives His research interests lie in sorting the neutral and adaptive processes that governed the diversification of populations

notes on contributors ix

and species during human evolution and in seeking the morpho logical correlates of these evolutionary processes using non‐invasive imaging techniques

Alan Brook is a member of the Craniofacial Biology Research Group in the School of Dentistry at the University of Adelaide Australia He also has an affiliation with Queen Mary University of London UK His main research interests revolve around studies of the human dentition aimed at clarifying the roles of genetic epigenetic and environ-mental factors on dental development and morphology

Scott E Burnett is Associate Professor of Anthropology at Eckerd College (St Petersburg Florida) His research concerns skeletal biology dental anthropology and bioarchaeology with emphasis on population variation He has published in anthropo-logical anatomical and clinical journals and a recent chapter ldquoWearrsquos the problem Examining the effect of dental wear on studies of crown morphologyrdquo (with JD Irish and MR Fong) in Anthropological Perspectives on Tooth Morphology Genetics Evolution Variation Scott and Irish eds (2013)

Charisse L Carver is a PhD candidate in the School of Human Evolution and Social Change at Arizona State University and a Fulbright Scholar Her research focuses on ethnogenesis and biodistance in the Early Middle Ages of Western Europe the bioar-chaeology of prehistoric Africa and dental morphology Recent publications include ldquoIncisor avulsion social identity and Saharan population history New data from the Early Holocene southern Saharardquo in Journal of Anthropological Archaeology

Frank P Cuozzo is an Associate Professor of Biological Anthropology at the University of North Dakota His research focuses on the variation and function of primate teeth with an emphasis on prosimian primates He has over a dozen years of field experience in Madagascar and South Africa where he studies the interaction between primate tooth morphology its use and the environments in which lemurs and other prosimians live thus studying their ldquodental ecologyrdquo

Lucas K Delezenersquos research focuses on the evolution of hominin and primate teeth His research has to date been primarily focused on canine and premolar evolution in early Australopithecus species It has ranged from investigating temporal trends in pre-molar evolution to reconstructing patterns of premolar and canine use using dental microwear analysis and to using patterns of trait covariation (morphological integration) to reveal the signatures of natural selection that drove early hominin dental evolution

Heather JH Edgar is Curator of Human Osteology Maxwell Museum of Anthropology and Assistant Professor of Anthropology University of New Mexico Her research focuses on how cultural and historical events and trends shape the biology of populations She often uses dental morphological data in this work these data can also be applied to race estimation Recent research includes biohistorical analyses of pre‐contact Mesoamerican groups and the development of theoretical and statistical approaches in forensic dental anthropology

Debbie Guatelli‐Steinberg is Professor of Anthropology and Courtesy Professor of Evolution Ecology and Organismal Biology at The Ohio State University Her research interests include developmental defects of enamel as well as dental growth development and morphology in living and fossil human and non‐human primates

x notes on contributors

Anna M Hardin is a PhD candidate in the Department of Anthropology at the University of Minnesota She is researching the impacts of body size and genetic heri-tability on primate dental morphology using a quantitative genetic approach Additional interests include fossil primate taxonomy and deciduous dental morphology

Edward F Harris is Professor Emeritus of Bioscience Research at the University of Tennessee Health Science Center He has published extensively in the areas of dental anthropology as well as dental clinical research He is also Editor Emeritus of the journal Dental Anthropology the official publication of the Dental Anthropology Association

Kelly Heim is currently working on her PhD in the Department of Anthropology at the University of Nevada Reno She received her MA from Louisiana State University Baton Rouge in 2013 where she also worked in the Forensic Anthropology and Computer Enhancement Services (FACES) Laboratory Her research interests include forensic anthropology dental development and dental morphology

Brian E Hemphill is an Associate Professor in the Department of Anthropology at the University of Alaska Fairbanks He has been actively conducting research on dental variability among prehistoric and living members of the myriad castes and tribes of South Asia since 1987 His primary research interests include multivariate statistical analyses and reconstruction of population histories especially with respect to patterns of gene flow genetic drift and marital distance

Simon Hillson is Professor of Bioarchaeology at the Institute of Archaeology University College London He has been teaching and researching in bioarchaeol-ogy since the completion of his PhD in 1979 Among his books published with Cambridge University Press are Teeth (first published 1986 second edition 2006) Dental Anthropology (1996) and Dental Development in Human Evolution and Bioarchaeology (2014)

Toby Hughes is a member of the Craniofacial Biology Research Group in the School of Dentistry at the University of Adelaide Australia His main research interests revolve around studies of the human dentition aimed at clarifying the roles of genetic epigenetic and environmental factors on dental development and morphology

Louise T Humphrey is a Researcher in Human Origins in the Department of Earth Sciences at the Natural History Museum in London Her research addresses human development and life history with a focus on retrieving the retrospective evidence of environmental and physiological influences embedded in the structure and chemical composition of dental tissues forming before and after birth

John P Hunter is Associate Professor of Evolution Ecology and Organismal Biology at The Ohio State University at Newark A vertebrate paleontologist he studies mam-mals of the late Mesozoic and early Cenozoic their evolution and ecology as revealed through their teeth and their early biogeographic history Recently in collaboration with Debbie Guatelli‐Steinberg and students he has turned his attention to one extant species of large mammal with particularly dull teeth and its relatives

notes on contributors xi

Joel D Irish is Professor of Bioarchaeology in the Research Centre in Evolutionary Anthropology and Palaeoecology at Liverpool John Moores University UK and Professor Emeritus in the Anthropology Department at the University of Alaska Fairbanks He is former Associate Editor of the American Journal of Physical Anthropology and his publications focus on dental nonmetric variation and biological affinity

Kent M Johnson is a PhD candidate in the School of Human Evolution and Social Change at Arizona State University His research involves state collapse and social orga-nization in the pre‐Hispanic Andes kinship peopling of the Americas and dental morphology Recent publications include ldquoSinodonty and beyondrdquo in Anthropological Perspectives on Tooth Morphology Genetics Evolution Variation and ldquoNew evidence on the spatiotemporal distribution and evolution of the Uto‐Aztecan premolarrdquo in American Journal of Physical Anthropology

Kristin L Krueger is an Assistant Professor of Anthropology at Loyola University Chicago She focuses her research on dietary and behavioral strategies of late Homo with particular emphasis on Neanderthals and recent modern humans Recent publica-tions include ldquoDental microwear texture analysis of hominins recovered by the Olduvai Landscape Paleoanthropology Project 1995ndash2007rdquo in Journal of Human Evolution and ldquoAnterior dental microwear texture analysis of the Krapina Neandertalsrdquo in Central European Journal of Geosciences

Adeline Le Cabec obtained her PhD in 2013 in the Department of Human Evolution at the MPI‐EVA from the Universities of Leipzig and Toulouse 3 She then investi-gated dental development in fossil hominins on the ID19 beamline at the ESRF and in the Department of Human Evolutionary Biology Harvard University She is currently a postdoctoral researcher at MPI‐EVA and on ID19 Her work focuses on developing new analytic techniques using synchrotron micro‐CT data

Clark Spencer Larsen is Distinguished Professor of Social and Behavioral Sciences and Chair of the Department of Anthropology at The Ohio State University in Columbus Ohio He received his PhD in biological anthropology from the University of Michigan Larsen is a Fellow of the American Association for the Advancement of Science former president of the American Association of Physical Anthropologists and Editor‐in‐Chief of the American Journal of Physical Anthropology His research focuses on biocultural adaptation during the last 10000 years of human evolution with particular emphasis on the history of health and lifestyle He currently directs or co‐directs field‐based research programs in Turkey Italy and the United States The study of teeth is central in all of his research and teaching Larsen is the author or editor of Advances in Dental Anthropology (co‐edited with Marc Kelley 1991) Bioarchaeology Interpreting Behavior from the Human Skeleton (2nd edn 2015) Skeletons in Our Closet Revealing Our Past through Bioarchaeology (2000) and the introductory textbook Our Origins Discovering Physical Anthropology (3rd edn 2014) He is the founding editor of the book series Bioarchaeological Interpretations of the Human Past Local Regional and Global Perspectives (University Press of Florida)

Loren R Lease is an Associate Professor in the Department of Sociology Anthropology and Gerontology at Youngstown State University Youngstown OH The focus of her research is on the morphology and metrics of the deciduous dentition

xii notes on contributors

Scott S Legge is an Associate Professor of Anthropology at Macalester College in St Paul MN His research focuses on human and non‐human primate skeletal pathology as well as non‐human primate dental variation and pathology His work has also included historic and prehistoric archaeology Additionally he is interested in growth and development nutrition and disease and human biology and adaptation

Helen M Liversidge is Reader in Dental Anthropology and a clinical teacher of pediatric dentistry at Barts and The Royal London School of Medicine and Dentistry Queen Mary University of London Her research interests include methods of dental age estimation from developing teeth and worldwide variation in tooth formation Key publications include The London Atlas of Tooth Development and Eruption reference data for deciduous teeth formation timing of third molar formation and interpreting dental maturity

Peter W Lucas is Professor of Bioclinical Sciences Faculty of Dentistry Kuwait University He holds a BSc from University College London and PhD and DSc from the University of London He worked at the National University of Singapore University of Hong Kong and George Washington University before joining Kuwait in 2011 He has about 120 full journal papers plus one book (Dental Functional Morphology 2004) His research revolves around feeding processes

Christopher Maier is a PhD student in the department of Anthropology at the University of Nevada Reno Previously he worked at the Forensic Anthropology and Computer Enhancement Services (FACES) Lab at Louisiana State University where he received his Masterrsquos degree His current research focuses on the use of cranial and dental morphology in the forensic assessment of ancestry and in broader questions of human variation His recent publications include ldquoPalate Shape and Depth A Shape‐Matching and Machine Learning Method for Estimating Ancestry from the Human Skeletonrdquo in Journal of Forensic Sciences

Mariacutea Martinoacuten‐Torres PhD in Medicine and Surgery MSc in Forensic Anthropology and MPhil in Human Origins is currently leading the Dental Research Group at the National Research Center on Human Evolution (Burgos Spain) As a member of the Atapuerca Research Team since 1998 she has research interests in hominin palaeobiol-ogy palaeopathology and the evolutionary scenario of the first Europeans She has led and participated in several international projects related to the study of hominin dental evidence worldwide

Greg C Nelson is an Adjunct in the Department of Anthropology at the University of Oregon Currently he is involved in the excavation and analysis of the Chelechol ra Orrak site Republic of Palau one of the earliest large cemeteries in Remote Oceania dating to c 3000 BP His research on these individuals focuses on determining their place of origin and understanding how these early settlers of the archipelago adapted to an isolated marginal environment

Kathleen S Paul is a PhD student and National Science Foundation Research Fellow in the School of Human Evolution and Social Change at Arizona State University Her research focuses on kinship deciduous dental morphology and the application of

notes on contributors xiii

multivariate dental data sets to genealogy reconstruction Her recent publications include ldquoBiological and spatial structure of an Early Classic cemetery at Charco Redondo Oaxacardquo in American Journal of Physical Anthropology

Anna LM Rautman is a PhD student at the University of New Mexico and the Chief Laboratory Assistant in the Maxwell Museumrsquos Laboratory of Human Osteology Her (2013) publication ldquoSecular Change in Dental Development in New Mexican Femalesrdquo in Dental Anthropology Journal described variation in the timing of dental development Her broader research interests include analyses comparing growth and development among different human body systems in order to better understand life history variables

Christopher W Schmidt received his PhD from Purdue University West Lafeyette IN in 1998 His research focus is on dietary reconstruction as indicated by dental microwear texture His ongoing DENTALWEAR project includes data from over 1000 people representing over 70 archaeological sites from around the globe Recent publications focus on the diets of people from Bronze Age England Canterbury Cathedral and Herculaneum His current focus is on understanding late Pleistocene and early Holocene foragers He is director of the UIndy Anthropology Graduate Program and Editor of Dental Anthropology

G Richard Scott is Professor of Anthropology at the University of Nevada Reno and Professor Emeritus at the University of Alaska Fairbanks where he taught for 24 years He has published widely on issues relating to tooth crown and root morphology and biological affinity

Christopher M Stojanowski is an Associate Professor in the School of Human Evolution and Social Change at Arizona State University He is the author of Bioarchaeology of Ethnogenesis in the Colonial Southeast and Mission Cemeteries and Mission Peoples Historical and Evolutionary Dimensions of Intracemetery Bioarchaeology both published by the University Press of Florida His research focuses on colonial populations of the Southeastern US and Early and Middle Holocene populations of the New World and Africa

Nancy Tang obtained her PhD in 2011 at the Institute of Archaeology University College London She is a postdoctoral fellow in the Exposure Biology Laboratory at the Icahn School of Medicine at Mount Sinai and a research associate in the Department of Human Evolutionary Biology at Harvard University She has a special interest in root dentine translucency (RDT) and is working on improving methods to study RDT in collaboration with the European Synchrotron Radiation Facility

Daniel H Temple is an Assistant Professor in the Department of Sociology and Anthropology at George Mason University Fairfax VA His research interests include developmental stress life history evolutionary morphology dental and skeletal biology diet and hunter‐gatherers His most recent publications address develop-mental stress life history trade‐offs the history of bioarchaeology and paleopatho-logical analyses among hunter‐gatherers from Northeast Asia These publications have appeared in the American Journal of Physical Anthropology PLoS One and Quarternary International

xiv notes on contributors

Grant Townsend is a member of the Craniofacial Biology Research Group in the School of Dentistry at the University of Adelaide Australia His main research interests revolve around studies of the human dentition aimed at clarifying the roles of genetic epigenetic and environmental factors on dental development and morphology

Peter S Ungar is Distinguished Professor and Chair of Anthropology at the University of Arkansas He is author of Mammal Teeth Origin Evolution and Diversity (2010) and Teeth A Very Short Introduction (2014) His research focuses on dental wear form and function He has published on a broad range of living and fossil animals from tyrannosaurids to modern humans

Robin Yong is a member of the Craniofacial Biology Research Group in the School of Dentistry at the University of Adelaide Australia His main research interests revolve around studies of the human dentition aimed at clarifying the roles of genetic epigenetic and environmental factors on dental development and morphology

Foreword

Clark Spencer Larsen

Kristin Krueger contributing author to this remarkable Companion volume (Chapter 24) says it best ldquoSimply put teeth are amazingrdquo All of us involved in the development of this book have long known this but by the time I got to Kruegerrsquos chapter her expres-sion of enthusiasm for the study of dentition in an anthropological perspective was clear for anyone reading beyond this foreword Teeth are amazing They are the only element of our skeleton coming into direct contact with the environment they are composed of dynamic tissues and they provide an incredible fund of data on so many fronts including physiology and stress adaptation evolution genetic variation social and cultural behavior diet and food use migration life history growth and development biological distance and relatedness anatomical variation oral and general health and identification and individuation in forensic contexts

From individual to population the dental record is central to understanding humans human‐like ancestors non‐human primates and mammals generally Teeth are dynamic they grow they develop and they have a nerve and blood supply That said though once formed the outer covering of teethmdashthe enamelmdashdoes not regenerate Encapsulated in each fully formed mature tooth is a permanent record unlike skeletal and all other tissues of the body As is so well articulated by Debbie Guatelli‐Steinberg in Chapter 27 the inability to regenerate lost or incompletely formed enamel is a problem for the individual in life but what a bonus for dental anthropologists to be able to document growth and other aspects recorded in the tooth

I regard this as the most important book on dental anthropology now available It contains something for anyone interested in teeth and at all levels from the beginning student to the senior scientist It successfully transcends the specialty foci by bringing the field into broad perspective via integration of the many lines and levels of inquiry ranging from morphology to ecology to diseasemdashall within the

xvi foreword

context of evolution the central and unifying theme that gives modern biological anthropology its identity and sense of mission

It is a thrill for me personally to see the record of advancement in dental anthropology so nicely encapsulated in this book It is all the more welcome since the last book of its kind Advances in Dental Anthropology (Kelley and Larsen 1991) was published 25 years ago The editors of Advances were especially interested in heralding the anthropological focus on teeth marking what a co‐editor of this Companion volume Richard Scott regards as the beginning of the ldquomaturationrdquo phase of the discipline It is breathtaking to see what has tran-spired in the last 25 years both in breadth and in scope Twenty‐five years ago I predicted extraordinary advances as the field moved forward but I would not have predicted the extent of production and the resources and quality of work that would bring the field to what it is today This new synthesis demonstrates a clear sense of passion accomplishment and mission in dental anthropology All 40 of the authors provide us with a new source a source that captures twenty‐first‐century developments theory and new ways of approach-ing the study of teeth A simple glance at the Companionrsquos table of contents reveals an authorship ranging from beginning dental anthropologists to senior leaders in the field a veritable whorsquos who of dental anthropologymdashthe experts and what they know best

While many of the chapters contain discussions of new quantitative methods and instrumentation that are opening a window onto the distant and recent past and the present what is especially remarkable are the advancements in hypothesis‐ and question‐driven science and the sense of where we need to go next to address central questions about the world of the primatemdashhuman and non‐human alike All of us in the field are amazed by the advances in methods and tools for addressing long‐standing problems For example the use of stable isotope analysis to track individual and population move-ments the development of ancient DNA analysis for identifying social groups and population relationships and the development of microwear and confocal analyses of occlusal surfaces of teeth for documenting tooth use and masticatory function It is also wonderful to see the improvements and revisions of the Arizona State University Dental Anthropology System (Turner Nichol and Scott 1991) for biodistance analysis of intra‐ and inter‐population affinities using crown and root traits Yet these areas of development and many more discussed in this Companion do not simply dote on the methodological accomplishments in the field Rather as the authors of the volume col-lectively demonstrate these accomplishments are viewed as tools to address long‐standing questions that our discipline has been asking and some for quite a few years What are the linkages between culture behavior and biology How have humans adapted to dramatic changes in their environments How can we identify and recon-struct social groupings in persons long deceased What is the role of the dentition in understanding health and living circumstances past and present What is the role of the dentition in mastication and how does it achieve critical functions relating to processing food as a beginning phase of digestion How best can we use such a complex process as dental development for understanding the myriad genetic epigenetic and environ-mental factors that determine morphology What is the relationship between genetics and development resulting in the shape size and number of teeth What are the key trends in primate and human dental evolution and how does the study of dentitionmdashsometimes the only recordmdashtell us about adaptation distribution and the extraordi-nary success over the course of the 50 million years or so of Order Primates What do sex and age differences in pathology size and variation mean in a human population undergoing fundamental changes in diet

foreword xvii

Several of the authors ask some quite critical questions about our sister disciplines in the life sciences In this regard Peter Lucas points out that the fundamental develop-ments in dental anthropology do not seem to play a central role in some of these disci-plines With regard to dental science Lucas suggests that the reason lies in the belief by some in the sciences that mechanical toothndashfood contacts are not especially important I believe that a central problem is the deficit of evolutionary and temporal context for both understanding dentition and dental function While participating in the instruction of orthodontics residents over the last couple of decades I am always struck by their relatively uninformed understanding of the context of occlusal and other abnormalities My contribution to an annual seminar on craniofacial growth and development for these residents has been to provide these future researchers andor practitioners with an overview of the evolution and ecology of human mastication The discussion of the evolution of mastication has provided the orthodontics residents with a wider perspec-tive on the integration of mastication dentition and craniofacial architecture

Read on If your interests in teeth and breadth of knowledge increase I will not be surprised

REFERENCES

Kelley MA and CS Larsen (eds) (1991) Advances in Dental Anthropology New York Wiley‐Liss

Turner CG II CR Nichol and GR Scott (1991) Scoring Procedures for Key Morphological Traits of the Permanent Dentition The Arizona State University Dental Anthropology System In MA Kelley and CS Larsen (eds) Advances in Dental Anthropology New York Wiley‐Liss pp 13ndash32

We wish to acknowledge mentors colleagues and friends recently lost First Christy G Turner II bioarchaeologist and dental anthropologist passed away in July 2013 Second Donald Morris bioarchaeologist and dental anthropologist passed away in August 2014 Christy was the chair and Don a member on both of our PhD commit-tees at Arizona State University To have one ldquotruerdquo dental anthropologist in an anthropology department is uncommon but to have two is exceptional Without them it is unlikely we would have been in a position even to attempt editing such a book as this Third Julius A Kieser passed away in June 2014 Jules as he preferred to be called focused on craniofacial biomechanics and forensic biology in the Faculty of Dentistry at the University of Otago New Zealand But before that he wrote Human Adult Odontometrics in 1990 for Cambridge University Press today it is a ldquomust readrdquo for dental anthropologists conducting dental metric study In fact Jules kindly agreed to write Chapter 19 for this volume Sadly it was not to be (Brian Hemphill kindly agreed to do both odontometric chapters)

We of course appreciate all of the hard work by the authors Obviously this book would not have been possible without them The folks at Wileymdashformerly and cur-rentlymdashhelped a lot including Rosalie M Robertson who shepherded this book through the process from proposal to acceptance From that point on Ben Thatcher Mark Graney and Elizabeth Swayze did the rest

Individually JDI thanks Clark Larsen for suggesting that I take on this project and for introducing me to Rosalie from Wiley Of course none of this would be possible without the faculty ldquoback in the dayrdquo at Arizona State University (PhD 1993) including Christy and Don as well as Charles Merbs Christopher Carr and Charles Redman among others and Minnesota State University (BS 1980 and MS 1984) including Richard Strachan Paul Brown and Michael Scullin who taught me about

Acknowledgments

acknowledgments xix

anthropology archaeology bioarchaeology and then dental anthropology It was a lot of work but also a lot of fun My parents provided me with lifelong support and my wife Carol has taken on this role for more than 20 years now

GRS owes a special thanks to Christy G Turner II and his entire family especially Korri and Olga in recent years for making me an unofficial family member Their unwavering support has allowed me to pursue the CGT II Legacy Project which will ultimately result in the release of materials (tables data sheets dental slides) that Christy accumulated during many decades across many continents In this way his enormous contribution to dental anthropology will live on for another generation After writing my first book with Christy my sons all pleaded with me never to write another book given the amount of time it took away from my fatherly duties Now they are all grown I doubt they mind if dad writes another book or two My three sons Garrett Geoffrey and Gunnar and three grandchildren Griffin Claire and Hadley remain an inspira-tion for dadgrandpa to keep pressing on

pART I ContextpART I

Chapter 1

A Companion to Dental Anthropology First Edition Edited by Joel D Irish and G Richard Scott copy 2016 John Wiley amp Sons Inc Published 2016 by John Wiley amp Sons Inc

Introduction to Dental Anthropology

Joel D Irish and G Richard Scott

The Foreword by Clark Larsen nicely addresses the content and purpose of this volume Beyond that the reader can look at the table of contents to ponder the topic of each chapter without us having to (re)state the obvious So we will keep this chapter short and to the point Think of it primarily as ldquoreading instructionsrdquo or at least suggested guidelines to get the most out of this book

What you hold before you whether tactilely or virtually is a textreaderreference book or if all else fails paperweight on the subject of all things teeth and tooth related That is we are talking teeth from the perspective of dental anthropologists So what is the definition of ldquodental anthropologyrdquo There is no need to reinvent the wheel when we have the internet According to the Medical Dictionary for the Dental Professions (2012) it is ldquo[a] branch of physical anthropology concerned with the origin evolution and development of dentition of primates especially humans and to the relationship between primatesrsquo dentition and their physical and social r elationshipsrdquo That d efinition works

The reasons why teeth are studied are numerous For one thing they are made up of the two hardest tissues in the body so are the most likely to be preserved in the fossil and archaeological records Think about it Many fossil primate species are defined based on teeth Given that teeth are the only components of the skeleton to come into direct contact with the environment we can learn about diet health and even certain cultural factors of individuals and groups Throw in the facts that tooth size and shape have high genetic components in expression and that unlike bone this expression does not remodel itself during life (other than via crown wear and pathology) and we have an excellent

ChApter 1

4 joel d irish and g richard scott

source to estimate biological origins and relationships across time and space There is much more but you can read about that in the remaining 30 chapters of this book

Who You Are And our SuggeStionS

At this point we would like to know something about you so we can offer suggestions on getting the most out of this volume Are you a dental anthropology beginner such as an upper‐division or graduate student taking an initial course in this ldquobranch of physical anthropologyrdquo or an interested layperson If so then you may be using this volume as a textbook or secondary class source In that case we suggest you read the rest of this chapter and then importantly skip directly to Chapter 7 The latter contains the fundamental terms and terminology needed to fully grasp the content of each remaining chapter Pay special attention to the differences in tooth class designations they vary among studies of primate fossil hominin and modern human dentitions (eg a lower first premolar may be labeled as P3 LP3 or LP1) After learning the basics flip back to Chapter 2 and read the rest of the volume in order

Are you an advanced graduate student post doc or a newly minted professional who wants to ldquobone uprdquo on dental anthropology or focus on specific topics of interest to your own research or teaching Then think of this book as a readerreference where you can reacquaint yourself with the basics and get up to speed on new methods areas of research and references andor gain some personal insight from experts in the field We suggest that you skim the rest of this chapter and start right in on Chapter 2 You should know enough to get by regarding dental terms and besides it will all come back to you as you peruse the material

Or are you a dental professional with strong academic or perhaps clinical experience who is inclined to use this volume for reference purposes If so then you can probably skip the rest of this chapter as well as Chapters 7ndash8 and perhaps any other chapter(s) covering material on which you are an expert Then go ahead and ldquocherry pickrdquo the sections and chapters that you find to be of interest

Volume orgAnizAtion And Content

This section heading may seem to imply that we are going to restate the obvious but as I have already said we are not Rather we are chiefly providing some rationale for the order and structure of chapters This book is organized exactly in the way the first editor has taught dental anthropology as an upper‐divisiongraduate‐level university course for the past couple of decades That is it proceeds in a more or less sequential manner both procedurally and temporally

part I Context (see Chapter 2) provides a diachronic review of where this subshydiscipline of physical (or biological) anthropology came from and a list of some of the major players involved

part II Dental evolution (Chapters 3ndash6) covers the origins and variation of teeth in an evolutionary perspectivemdashfrom their first appearance in non‐primates (which according to the above definition is actually outside the realm of dental anthropology) to variation among non‐human primates and fossil hominins including pre‐modern Homo sapiens

introduction to dental anthropology 5

part III the human Dentition (Chapters 7ndash9) introduces the terminology a natomical structures and functions of the teeth and supporting structures that are necessary for dental anthropological research

part IV Dental Growth and Development (Chapters 10ndash13) provides key information on dental genetics crown and root formation eruption final form and variation therein

part V Dental histology from the Inside Out (Chapters 14ndash16) continues on from the previous part by providing detail on the structure and material properties of tissues that comprise the teeth

part VI Dental Morphometric Variation in populations (Chapters 17ndash20) shows how tooth shape and size can be quantified in samples to estimate intra‐ and inter‐population variation and affinities that is involving the ldquobig picturerdquo in anthropology

part VII Dental Morphometric Variation in Individuals (Chapters 21ndash24) follows up on the preceding part by focusing on individual characterization and d ifferences forensic applications are essential but the ability to reconstruct life h istories as in an archaeological context is necessary as well

part VIII Dental health and Disease (Chapters 25ndash28) covers the importance of dental pathology like caries and enamel hypoplasia for understanding life histories (including diet indications of stress etc) at both the population and individual levels

part IX the Future of Dental Anthropology (Chapters 29ndash31) shows a glimpse of where the subdiscipline is headed by describing state‐of‐the‐s cience approaches to assess the link among morphological variants the use of chemical analyses and an overview of non‐destructive techniques to image the inside of teeth All in all the future looks bright

expertiSe And A perSonAl touCh

No matter whether you use this book as a text reader reference or something else it is important to know that each chapter was written by an expert (or experts) in that area of dental research For example do you want to know about the origins and early evolution of teeth The author of Chapter 3 Peter Ungar (2010) literally wrote the book on the subject The same goes for Peter Lucas (2004) concerning jaw function in Chapter 9 and the authors of Chapter 16 on enamel structure in the latter case Daniel Antoine (2001) wrote his entire PhD thesis on enamel() and Simon Hillson (1996 2000 2014) has no fewer than three books in which this tissue is detailed However a quick perusal of the references in these chapters shows that the authors do much more than write books as the many publications in peer‐reviewed professional journals i ndicate Along these lines check out all the articles by the first author of Chapter 13 on tooth classes and the field concept Grant Townsend (eg among others 2005 2009) and Chapter 27rsquos author Debbie Guatelli‐Steinberg (eg among others 2004 2012) on dental stress indicators (enamel hypoplasia) Each of the remaining authors could be singled out for kudos in this manner but there are just too many So please make sure to look through their short biographies at the beginning of this book to get an idea of who they are

Now think about this If you were tasked with creating a textbook (or readerr eference) on any topic of your choosing would you not want it to be written by the best and the brightest Ordinarily textbooks are written by one or a few authors Such

6 joel d irish and g richard scott

an approach is great for chapter consistency in writing style format and so forth Nevertheless those few authors are likely not experts on every topic in the book For example wemdashyour kindly book editorsmdashare dental morphologists and we also know a little about other aspects of dental anthropology However we are not experts in p rimatology paleoanthropology genetics forensics or histology So we got the experts to write about what they know best while doing a ldquolight touchrdquo editing job to strive for consistency across chapters in writing style format and so forth But wait there is more Beyond presenting material specific to each chapter topic many authors provided findings from their own research maybe a case study or two andor voiced opinions about their or other areas of dental anthropology As such they p ersonalized their chapters so you can see that there is more to research than just the same old ldquoempiricalrdquo or ldquoscientificrdquo approach

noW on to ChApter 2 (or ChApter 7)

So if you have read this far you must not be a dental expert unless you simply wanted to see how everything turned out in the end In any event sit back tactilely or virtually grab this volume firmly and start reading (and above all learning) The 40+ authors put a lot of work into their chapters We hope you enjoy reading them as much as we did when putting this edited volume together

reFereNCeS

Antoine D (2001) Evaluating the Periodicity of Incremental Structures in Dental Enamel as a Means of Studying Growth in Children from Past Human Population PhD thesis University College London

Guatelli‐Steinberg D CS Larsen and DL Hutchinson (2004) Prevalence and the Duration of Linear Enamel Hypoplasia A Comparative Study of Neandertals and Inuit Foragers Journal of Human Evolution 47 65ndash84

Guatelli‐Steinberg D RJ Ferrell and J Spence (2012) Linear Enamel Hypoplasia as an Indicator of Physiological Stress in Great Apes Reviewing the Evidence in Light of Enamel Growth Variation American Journal of Physical Anthropology 148 191ndash204

Hillson S (1996) Dental Anthropology Cambridge Cambridge University PressHillson S (2000) Teeth 2nd edition Cambridge Cambridge University PressHillson S (2014) Tooth Development in Human Evolution and Bioarchaeology Cambridge

Cambridge University PressLucas PW (2004) Dental Functional Morphology How Teeth Work Cambridge Cambridge

University PressMedical Dictionary for the Dental Professions (2012) Dental Anthropology Retrieved December

7 2014 from httpmedical‐dictionarythefreedictionarycomdental+anthropologyTownsend G Harris EF Lesot H Clauss F and Brook AH (2009) Morphogenetic

Fields within the Human Dentition A New Clinically Relevant Synthesis of an Old Concept Archives of Oral Biology 54S S34ndashS44

Townsend G Richards L Hughes T Pinkerton S and Schwerdt W (2005) Epigenetic Influences May Explain Dental Differences in Monozygotic Twin Pairs Australian Dental Journal 50 95ndash100

Ungar PS (2010) Mammal Teeth Origin Evolution and Diversity Baltimore MD Johns Hopkins University Press

Chapter 1

A Companion to Dental Anthropology First Edition Edited by Joel D Irish and G Richard Scott copy 2016 John Wiley amp Sons Inc Published 2016 by John Wiley amp Sons Inc

A Brief History of Dental Anthropology

G Richard Scott

Several papers have been written on the history of dental anthropology from both world (Dahlberg 1991 Scott 1997 Scott and Turner 2008) and regional (Brown 1992 1998 Kosa 1993) perspectives To avoid duplication this brief history adopts a different approach It brackets developments in three periods foundationsmdashthe period preceding the classic edited volume Dental Anthropology (Brothwell 1963) developmentmdashfrom 1963 to Advances in Dental Anthropology (Kelley and Larsen 1991) and maturationmdashfrom 1991 to the present The emphasis is on the various ways anthropologists have used teeth to further our understanding of primate and human evolution variation health and behavior

Foundations (nineteenth Century to 1963)

The role of the physical or biological anthropologist is to describe biological variation and explain it in terms of adaptation evolution and history In pre‐Darwinian times anthroshypologists focused largely on racial variation and classification Teeth played no role in the early classifications of Blumenbach Cuvier and others who focused on externally visible physical characteristics like skin hair and eye color and form and skull types

In the nineteenth century a handful of anthropologists started exploring the utility of teeth and their role in understanding human health and behavior (eg P Broca and crown wear LH Mummery and oral pathology) and variation (WH Flower and tooth size) More substantive developments were taking place in paleontology where

CHApter 2

8 g richard scott

Richard Owen Edward Drinker Cope and Henry Fairfield Osborn were laying the foundations of comparative odontology (Peyer 1968)

With the post‐Darwinian acceptance that humans were primates and their closest relationship was to apes more researchers started paying heed to tooth size and m orphology in primates the still limited array of fossil hominins and recent human populations For a broad‐scale approach a major contribution was The Origin and Evolution of the Human Dentition by William King Gregory (1922) Although Gregory discussed tooth morphology in humans he minimized the significance of variation among the geographic races mostly because he was familiar with a limited number of variables like upper and lower molar cusp number Carabellirsquos trait and shovel‐shaped incisors Around this time Hrdlička (1920) wrote the classic paper on shovel‐shaped teeth wherein he noted the close similarity between Native Americans and north Asians and their contrast to European and African populations

Hrdlička was among the first researchers to appreciate the extent of dental variation among world populations because he had access to thousands of skeletal remains at the US National Museum of Natural History Other significant contributions in this period were regional in scope Classic examples include the characterization of the Australian Aboriginal dentition by TD Campbell (1925) and the Bantu by JC Middleton Shaw (1931) Slightly later PO Pedersen (1949) and CFA Moorrees (1957) produced classic monographs on the dentitions of East Greenland Eskimos and Aleuts r espectively These various authors as well as the many referenced below will be cited throughout this volume

The monographs already noted plus significant journal articles on the Indians of Pecos Pueblo (Nelson 1938) and Texas Indians (Goldstein 1948) provided descripshytions of tooth size morphology dental pathologies and crown wear Comparative data were still limited During this period a seminal paper by RW Leigh (1925) focused on the contrasts in dental health between tribes who lived in different environments and practiced diverse subsistence activities Leigh was among the first to demonstrate clearly the impact of agriculture (and agricultural practices such as grinding grain) on the human dentition

Some researchers including AA Dahlberg (Figure 21a) focused primarily on tooth morphology One of his most important early contributions was ldquoThe Changing Dentition of Manrdquo (Dahlberg 1945) in which he laid out the fundamental principles of morphogenetic fields in the human dentition as a follow‐up to the ldquofield theoryrdquo papers of PM Butler (1937 1939) Following the modern evolutionary synthesis of the early 1940s anthropologists started thinking more in terms of genes than types stimulating papers such as ldquoGenetic Analysis of Racial Traits of the Teethrdquo (Lasker 1950) and subsequent attempts to decipher the modes of inheritance of common morphological traits (cf Kraus 1951 1957 Kraus Wise and Frei 1959 Lasker 1957)

By the mid‐1950s AA Dahlberg (1956) determined that the systematic study of human dental variation was impeded by a lack of standardization Toward that end he developed a series of plaster plaques that showed ranked expressions for key morphoshylogical traits of the human dentition including shovel‐shaped incisors the hypocone Carabellirsquos trait and the protostylid These were widely adopted and observations based on these standards appeared in many articles for the next two decades

Prior to 1963 studies of fossil hominin teeth were mostly descriptive in nature Two key monographs did appear during this time ldquoThe Dentition of Sinanthropus pekinensisrdquo (Weidenreich 1937) and The Dentition of the Australopithecinae (Robinson 1956)