BIOLOGICAL ANTHROPOLOGYOF THE HUMAN SKELETON

Second Edition

Edited by

M. ANNE KATZENBERGDepartment of Archaeology, University of Calgary

SHELLEY R. SAUNDERSDepartment of Anthropology, McMaster University

Innodata9780470245835.jpg

BIOLOGICAL ANTHROPOLOGYOF THE HUMAN SKELETON

BIOLOGICAL ANTHROPOLOGYOF THE HUMAN SKELETON

Second Edition

Edited by

M. ANNE KATZENBERGDepartment of Archaeology, University of Calgary

SHELLEY R. SAUNDERSDepartment of Anthropology, McMaster University

Copyright# 2008 by John Wiley & Sons, Inc. All rights reserved

Published by John Wiley & Sons, Inc., Hoboken, New JerseyPublished simultaneously in Canada

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To our families

to Steve and MartyM. Anne Katzenberg

to Victor, Rob, and BarbShelley R. Saunders

CONTENTS

PREFACE TO THE SECOND EDITION—M. ANNE KATZENBERGAND SHELLEY R. SAUNDERS xi

PREFACE TO THE FIRST EDITION—M. ANNE KATZENBERGAND SHELLEY R. SAUNDERS xv

ACKNOWLEDGMENTS xxi

CONTRIBUTORS xxiii

FOREWORD—JANE E. BUIKSTRA xxxiii

PART I THEORY AND APPLICATION IN STUDIES OF PAST PEOPLES 1

1 Bioarchaeological Ethics: A Historical Perspective on theValue of Human Remains 3Phillip L. Walker

2 Forensic Anthropology: Methodology and Diversity ofApplications 41Douglas H. Ubelaker

3 Taphonomy and the Nature of Archaeological Assemblages 71Ann L. W. Stodder

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PART II MORPHOLOGICAL AND DEVELOPMENTAL ANALYSES 115

4 Juvenile Skeletons and Growth-Related Studies 117Shelley R. Saunders

5 Histomorphometry of Human Cortical Bone: Applications toAge Estimation 149Alexander G. Robling and Sam D. Stout

6 Biomechanical Analyses of Archaeological Human Skeletons 183Christopher B. Ruff

7 Morphometrics and Biological Anthropology in the Postgenomic Age 207Benedikt Hallgr|¤msson, Miriam Leah Zelditch, Trish E. Parsons,Erika Kristensen, Nathan M. Young, and Steven K. Boyd

8 Reading Between the Lines: Dental Development andSubadult Age Assessment Using the MicrostructuralGrowth Markers of Teeth 237Charles M. FitzGerald and Jerome C. Rose

9 Dental Morphology 265G. Richard Scott

PART III PREHISTORIC HEALTH AND DISEASE 299

10 Dental Pathology 301Simon Hillson

11 Analysis and Interpretation of Skeletal Trauma 341Nancy C. Lovell

12 Light and Broken Bones: Examining and InterpretingBone Loss and Osteoporosis in Past Populations 387Sabrina C. Agarwal

PART IV CHEMICAL AND GENETIC ANALYSES OF HARD TISSUES 411

13 Stable Isotope Analysis: A Tool for Studying Past Diet,Demography, and Life History 413M. Anne Katzenberg

viii CONTENTS

14 Bone Chemistry and Trace Element Analysis 443James Burton

15 DNA Analysis of Archaeological Remains 461Anne C. Stone

PART V QUANTITATIVE METHODS AND POPULATION STUDIES 485

16 Metric Analysis of Skeletal Remains: Methods and Applications 487Michael Pietrusewsky

17 Nonmetric Trait Variation in the Skeleton: Abnormalities,Anomalies, and Atavisms 533Shelley R. Saunders and Dori L. Rainey

18 Advances in Paleodemography 561George R. Milner, James W. Wood, and Jesper L. Boldsen

19 Method and Theory in Paleodemography, with anApplication to a Hunting, Fishing and GatheringVillage from the Late Eastern Woodlands of North America 601Richard S. Meindl, Robert P. Mensforth, and C. Owen Lovejoy

Index 619

CONTENTS ix

PREFACE TO THE SECOND EDITION

The preface to the first edition sets out thegoals we hoped to accomplish by preparingthe volume titled, Biological Anthropology ofthe Human Skeleton. In this second editionwe endeavor to maintain those goals, to updaterecent developments in skeletal studies andparticularly, to emphasize information that pro-vides the reader with a basic understanding ofthe various techniques and methods of investi-gating bones and teeth. Many chapters includeexamples, set off from the main body of thetext, that illustrate or offer more detail aboutthe particular analysis under consideration. Wealso provide six new chapters on topics notcovered in the previous edition. These topicsinclude taphonomic factors affecting burialassemblages, nonmetric traits of the skeletonand dentition, trauma, osteoporosis, and newdevelopments in morphometric analysis. It isour hope that the book will be used in upperlevel undergraduate and graduate courses inhuman skeletal studies (e.g., advanced humanosteology) as well as by interested professionalsseeking a better understanding of advancedmethods in osteological research. The chaptersshould provide an entry point into a particularspecialty, with background information as well

as practical guidelines, applications, and criticalreviews of research approaches, including awealth of selected references for additionalreading.

The book is divided into five parts,although considerable overlap exists and somechapters could have easily appeared in one oranother section. Part I is titled “Theory andApplication in Studies of Past People” andincludes three chapters. As before, the firstchapter, on ethical considerations of workingwith human skeletal remains, is presentedby Phillip Walker. Walker has updated andexpanded the scope of his chapter to includeworldwide examples of problems and solutionsto working with human remains. The secondchapter, by Douglas Ubelaker, provides currentperspectives on the interrelationship betweenforensic anthropology and more traditionalstudies in human osteology, arguing that theseare complementary fields of inquiry. The fieldof forensic anthropology has gained promi-nence since the publication of the first editionof this book, with an increase in the number ofacademic positions in the field, and increasedparticipation by forensic anthropologists inmedico-legal investigations. The third chapter

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in the first section is a new contribution by AnnL. Stodder, on taphonomy and human skeletalremains. Stodder draws from her experience inworking with burials and specific burial con-texts in several different regions of the worldto offer a comprehensive review of the variouspostmortem factors that affect the integrity ofthe skeleton after death.

Part II is newly titled “Morphologicaland Developmental Analyses” and includesfive chapters on development and modelingof bones and teeth. “Juvenile Skeletons andGrowth Related Studies,” by ShelleySaunders, examines the problems of studyingjuvenile skeletal remains from archaeologicalsites. This chapter has been updated with newexamples of applications from Saunders’ exten-sive work with historic cemeteries. AlexanderRobling and Sam Stout have revised theirprevious contribution on histomorphometryand, once again, provide helpful appendices,including a worked example of age deter-mination from cortical bone histology and acompilation of various histomorphometrictechniques for age determination from variousskeletal elements. Christopher Ruff has updatedhis chapter on biomechanical analyses, provid-ing new examples and illustrations. BenediktHallgrimsson and colleagues present a newchapter on the “new morphometrics” and theimportance of understanding the interfacebetween morphometric studies in the biologicalsciences and those studies in biological anthro-pology in the context of amore solid understand-ing of genetic mechanisms and their role indetermining phenotypic variation. The chapteron dental microstructure, by Charles FitzGeraldand Jerome Rose, retains the clear descriptionsof the microscopic structure of teeth and theevents recorded indentalmicrostructures, includ-ing evidence for stress and for age determination.Examples of more recent applications and newtechnological developments have been added.The final chapter in this section is a new contri-bution by Richard Scott on dental morphology,specifically dental nonmetric traits. The chapterprovides descriptions of various morphological

variants of the teeth and very practical adviceon how to recognize and record dental crowntraits so that they can be used in populationstudies.

Part III, “Prehistoric Health and Disease,”includes three chapters. Simon Hillson hasupdated his previous chapter on dental patho-logy, retaining a protocol for data collectionand updating the state of our understanding ofthe causes and implications of pathological con-ditions of the teeth and supporting structures.Nancy Lovell provides a new chapter on skeletaltrauma. This chapter focuses on fractures butalso includes more general information onresponses of bone to trauma and diagnosticprocedures for evaluating trauma in the past.The third chapter is also new to the secondedition. Sabrina Agarwal presents informationon osteoporosis in past populations with bothcross-cultural and historical perspectives.She offers a very useful comparison of theadvantages and disadvantages of the differentmethodological techniques for obtaining infor-mation on bone mass, density, and quality andtheir relevance to the study of past populations.

Part IV, “Chemical and Genetic Analysesof Hard Tissues,” as in the previous edition,includes three chapters. Anne Katzenbergdescribes methods and applications of stableisotope analysis that are used to reconstructdiet, estimate the duration of nursing, and deter-mine residence and migration patterns of thepast. This field has expanded considerablysince the previous edition. James Burton dis-cusses bone chemistry and the trace elementsof bone that have been used to reconstruct pastdiet as well as studies focusing on postmortemalterations of bone chemistry. Anne Stone pro-vides background and examples of ancientDNA studies from human remains. She illus-trates the significant challenges of workingwith ancient DNA, the fact that it is highlysubject to destruction and contamination andexpensive to analyse. She also points out thathypotheses about the genetics of populationsin the past must be consistent with what isknown about modern populations and that

xii PREFACE TO THE SECOND EDITION

research projects require the coordination ofexpertise from many different fields frompaleopathology to archaeology.

Part V includes four chapters on quantita-tive methods and population studies. MichaelPietrusewsky has updated his contributionon metric analysis, focusing on craniometricstudies for population reconstruction. He pro-vides an example from his extensive work inPolynesia. Shelley Saunders and Dori Raineyprovide a new chapter on skeletal non-metrictraits. They critically review the background ofsuch studies and include illustrations of manyof the more commonly used traits. They offersuggestions for future areas of research in thisfield, including study of the ontogenic develop-ment of specific traits and the relationshipbetween the prevalence of traits in past popu-lations and information on other skeletal criteria

such as DNA studies. As in the previous edition,there are two chapters on paleodemography.George Milner, James Wood, and JesperBoldsen have updated their previous contribu-tion focusing on both the promises and thelimitations of paleodemographic studies. A newchapter by Richard Meindl, Robert Mensforth,and Owen Lovejoy provides a detailed exampleof a particular paleodemographic study, fromthe Libben site, in northern Ohio.

As with the previous edition, we hope thatthis volume will provide useful informationfor both current and future biological anthropol-ogists interested in the latest research on humanskeletal and dental remains.

M. ANNE KATZENBERGSHELLEY R. SAUNDERS

PREFACE TO THE SECOND EDITION xiii

PREFACE TO THE FIRST EDITION

What’s Bred in the Bone, a novel by RobertsonDavies, begins with the proverb, “What’s bredin the bone will not out in the flesh.” The storyis about a man who supposedly reflects his“breeding” since his behavior and character-istics are direct reflections of what he has inher-ited from his family. Although biologicaldeterminism may work in fiction, it is anathemato the biological anthropologist. The corner-stone of biological anthropology is the inter-action of culture and human biology. What ismanifested in the physical and behavioralcharacteristics of any living being is a result ofthe intertwining of an inherited genome withenvironmental factors. Human osteologistshave struggled with this concept from the earl-iest beginnings of skeletal studies and continueto struggle with it today. Ancient DNA studiessuggest that we ultimately want to knowthe “inherent” properties coming out of thebones. If we could read the genome, we would“know” the person. But of course, we under-stand that, as living tissues, bones and teethare influenced by environmental forces. Bonesrespond to mechanical forces, and thus, theyalter in response to activities and stresses.Craniometric studies attempt to study popu-lation relationships, assuming that cranialshape and size reflect inherited features, but

we know that cranial shape and size can bealtered purposefully (head binding) or uninten-tionally (chewing stresses). It is the job of thehuman osteologist to study the interactionsbetween inherited characteristics and theirmodification by the environment in order tounderstand, not just what is “bred” in the bonebut also what bones can tell us about the flesh,that is, the lives of earlier peoples.

Each of the following chapters deals with aspecific type of advanced analysis of bones andteeth. The original plan for the book was to bea second edition of our earlier edited book,Skeletal Biology of Past People: ResearchMethods. However, as work progressed, itseemed that with five additional chapters andmany new contributors, it is really somethingdifferent. The differences are directly relatedto changes that have occurred in the analysisof human skeletal and dental remains over thepast few years. Most notably these changesinclude heightened ethical concerns aboutstudying the skeletal remains of aboriginalpeoples in many countries where those peopleare no longer the dominant culture. These con-cerns and the resulting legislation in some juris-dictions have radically changed the wayphysical anthropologists and archaeologistscarry out their work. A second change is the

xv

rise of forensic anthropology and the fact thatresearch in forensic anthropology, while stilloverlapping with more traditional approaches,now includes topics not central to studies ofarchaeological skeletons. We begin this bookwith chapters on the ethics of studying humanremains and forensic anthropology.

An important theme that is found throughoutthe book is the progress of new methods. Wewere training to become anthropologists inthe 1970s when many new research areas wereemerging in physical anthropology. The earlierpractice of providing descriptive osteologicalreports either as stand-alone works or, morecommonly, as appendices to archaeologicalsite reports was fading out and more problem-oriented research was emerging. Biologicaldistance studies using both metric and non-metric traits on human bones and teeth werecarried out in order to investigate prehistoricmigration and relatedness through time andspace. Paleopathology was emerging as ameans of addressing questions about pre-historic adaptations in contrast to the earlieremphasis on unusual cases of specific diseases.Paleodemography, similarly, addressed ques-tions of adaptation of earlier populations.Since the initial enthusiastic studies all ofthese topics have undergone criticism and haveemerged as, perhaps, humbled, but also strength-ened, by the critiques. The same is true of themore recently introduced methods involvingbiochemical analyses of bones and teeth. Thesemethods include analyses of trace elements,stable isotopes, and ancient DNA.

Each of these methods has undergone aseries of stages that may be characterized asfollows:

† Discovery—either entirely new or new tophysical anthropology, a new method isdiscovered and the potential applicationsare explored.

† Applications to questions of interestregarding reconstructing past peoples.

† Critique, introspection, experimentation.† Emergence in a stronger, more reasonedform.

NAGPRA (Native American GravesProtection and Repatriation Act) and similarlegislation in other countries have led to a recon-figuring of how skeletal studies of past peoplesare carried out. Some of these changes can beviewed in a positive light. For example, stan-dards have been developed in the expectationthat collections will not be curated indefinitely.These standards were needed even before theprospect of reburial emerged. In addition, aninteresting configuration of events happened inthe 1990s. As some Native Americans voicedtheir disapproval of skeletal studies, expandingurban development led to archaeological exca-vations of several large, historic cemeteriesdating from the eighteenth and nineteenth cen-turies. These cemeteries contained the remainsof Euro-americans and African Americans aswell as other groups. At the same time, thegrowing number of students trained in humanosteology provided a pool of individuals toexcavate and study these remains. Debatesabout excavation and study continued but inmany cases some period of time was allowedfor proper scientific study. One specialexample of the cooperation between scientistsand concerned descendants is the work beingconducted at Howard University on a largeAfrican-American slave cemetery discoveredin New York City. In Europe, there is a longhistory of excavating historic cemeteriesand the increasing number of trained humanosteologists has led to larger scale studies(the St. Brides’ skeletal collection in London,England is a good example). The increasedscientific study of skeletons from historic ceme-teries has also provided opportunities for testingmethods. In many cases, the identities of at leastsome individuals are known from legible coffininscriptions or detailed cemetery maps. It hasbeen possible to investigate the accuracy ofmethods of determining sex and age at deathand to detect biases in mortality samples thatare directly related to causes of death.

This book is organized into five parts. Part I,theory and application, features two chaptersthat describe recent shifts in skeletal studies.Walker’s chapter provides information on how

xvi PREFACE TO THE FIRST EDITION

humans have regarded the dead over time andacross cultures. He grapples with the issuessurrounding the ethics of skeletal research, theclash with cultural beliefs about treatment ofthe dead, and the politics of communities.Taking a clearly anthropological approach tothese questions, he shows us that there is a tre-mendous diversity of attitudes about the phys-ical remains of the dead. He makes a strongcase for the value of and the justification of scien-tific research. Ubelaker focuses on the develop-ment of forensic anthropology with its roots indescriptive osteology and its current form asan applied specialization of human osteology.He discusses the major comparative collectionsused for establishing standards, includingthe recently developed forensic data bank.He then takes the reader through the varioussteps in forensic anthropology, includingrecovery, identification, sex and age determi-nation, stature estimation, and positive identifi-cation. He concludes with information ontraining opportunities and professional organiz-ations dedicated to forensic anthropology.

Part II includes chapters on morphologicalanalyses of bones and teeth and age changes.Four of these contributors prepared chaptersfor our earlier book, and although the topicsare similar, each chapter includes contributionsand advances that have occurred throughout the1990s. Ruff describes biomechanical analysesof bones and the applications of such studiesto understanding past human behavior rangingfrom fossil hominids through to early historichuman groups. He draws from his ownextensive research to provide examples ofhow biomechanical studies have improvedour understanding of past activity patterns.Examples include changes in robusticitythroughout human evolution, the relationshipbetween subsistence and bone strength, andthe relationship between gender roles and theirbiological manifestation in bone structure.Mayhall covers dentalmorphology highlightingnewer methods of characterizing tooth size andshape, and the applications of such studies tobiological and behavioral characteristics ofpast peoples. He emphasizes the importance of

achieving precision of observations of bothdental measures and dental morphologicaltraits. He also argues for maintaining simplicityin our methodological approaches. Both ofthese aspects of the research process are absol-utely necessary for us to make meaningful com-parisons of the results obtained by differentobservers. Mayhall shows that knowledge inthe field of dental morphology remains limitedbecause the precision necessary for properlyevaluating population variability has still notbeen achieved. Saunders covers the varioustypes of studies that are specific to subadults,focusing on age determination but also consid-ering sex determination and variations ingrowth and development. One problem withproceeding to studies of growth and develop-ment is that of sampling. Differential burialpractices, differential preservation, and biasesrelated to cause of death can all cause problemsin assessing past growth patterns from subadultburials. Some of these problems have beenaddressed in studies of a large historic cemeterywhere parish records are available for compari-son. This cemetery has also provided opportu-nities for assessing historic variation in growthand development as well as for testingmethods of age determination. Saunders andher students have demonstrated how carefulstudy of historic samples can not only tell usmore about those particular people but alsocan help us to evaluate methods used on prehis-toric samples. FitzGerald and Rose presentinformation on age determination for subadultremains through dental microstructure analysis.The use of newer image analysis techniques(which are noweasy to install inmost anthropol-ogy laboratories) improves precision andrelieves the tedium of collecting these data.This research shows great promise. If we canget a clearer picture of the amount of inter-and intrapopulation variation in dental develop-ment, we will know more about how tissuegrowth is buffered from stress and whethermeaningful population differences really doexist. As these authors explain, it is only veryrecently that the investigation of microstructuralgrowth markers in dental tissues has become

PREFACE TO THE FIRST EDITION xvii

accepted as appropriate for estimating toothcrown formation times. Robling and Stoutprovide details as well as examples of adultage determination based on bone histomorpho-metry. They review the principles of bone mod-eling and remodeling as a prelude to explaininghow cortical bone microstructure is used in agedetermination. Variations caused by activity,sex, disease, and population affinity are dis-cussed. Appendices to their chapter allow oneto practice the methods of histological agedetermination on photomicrographs from afemur and a rib.

Part III is titled “Prehistoric Health andDisease” and includes three chapters. As inPart II, the sequence of chapters is as follows:studies based on gross observations of bones,gross observations of teeth, and microscopicstudies. Lovell focuses on paleopathology anddiagnosis of bony lesions. She provides detailedinformation on various diagnostic methods,including radiology and microscopy. Stepstoward diagnosis are discussed with emphasison accurate description and consideration ofthe distribution of lesions within an individualskeleton as well as the distribution withinskeletal samples. Hillson presents methods foranalyzing and describing dental pathology,with detailed information on the underlyingcauses of various conditions. He stresses theimportance of careful observation, demonstrat-ing how different ways of scoring pathologicalchanges can dramatically alter determinationsof disease prevalence. If care is taken withobservations, so that the surviving jaws andteeth in skeletal collections really do representwhat was buried, then the distribution ofdental disease can tell us a lot about the dietsand activities of past populations. Then we canseek correspondence between dental data anddata from stable isotopes, faunal and botanicalassemblages, and artifacts used in daily life.Pfeiffer covers the subject of bone histologywith respect to healthy bone turnover andvarious disease states. This chapter ties innicely with those of Ruff and Robling andStout in that it covers information on bone struc-ture at the histological level and the factors

that account for variation. Her work includesvariation in bone histology over recent humanevolution with examples drawn fromNeandertals to recent European immigrants toCanada. Procedures for preparing bones inthin sections are reviewed with cautions regard-ing diagenetic alteration.

Part IV, “Chemical and Genetic Analysesof Hard Tissues,” includes chapters on stableisotope analysis, trace element analysis, andancient DNA. Katzenberg provides backgroundinformation on stable isotope studies andexamples of applications to questions regardingpaleodiet, migration, and life history. Shedemonstrates how isotopic analysis of archaeo-logical tissues has advanced dramatically over arelatively short time span. Rather than simplyconfirming information that was already avail-able from other sources, she shows how thisfield has called into question various archaeolo-gically hypotheses about subsistence adapta-tions as well as adding to our understanding ofhuman ecology. She discusses three areas ofresearch that are particularly promisingbecause of their implications for a more detailedreading of the past. These areas include recon-structing infant feeding practices, detectingpathological changes in bones, and the manage-ment of animal and plant species by earlierhuman populations. Sandford and Weaver pro-vide information on the current status of traceelement studies. These studies include attemptsto control for postmortem changes. They focustheir discussion on the dietary indicators, stron-tium and barium, and the toxic element, lead.This chapter nicely illustrates the stages ofnew methods, discussed early in the preface.Sandford and Weaver have labeled these“Inaugural” (discovery and early applications),“Intermediate” (reevaluation and testing), and“Modern” (emphasis on experimental andsimulation studies). The chapters on stableisotope analysis and trace element analysisboth emphasize the importance of trainingin the physical sciences. Stone discussesadvances in the isolation and analysis ofancient DNA. A great wave of excitement wasushered in with the first developments in the

xviii PREFACE TO THE FIRST EDITION

extraction and amplification of ancient DNA.If we can retrieve fragments of genes fromlong deceased humans, surely we canreconstruct the evolutionary and populationhistory of past human groups. But the earlyclaims for the retrieval of ancient DNA fromdinosaurs and other fossils were cast asidewhen it was shown that the amplified DNAcame from modern contaminants. The promiseof ancient DNA research lost some of itsluster. Yet, more recently, Stone was part ofthe research team able to offer clear evidencefor the sequencing of Neanderthal DNA.Nevertheless, she cautions us about the difficul-ties of proving positive results and warns usthat the promise is there, but the road ahead isstill difficult.

Part V, “Quantitative Methods andPopulation Studies,” includes three chapters.Pietrusewsky discusses metric techniquesand their applications to biological distancestudies. He takes the reader through thevarious statistical procedures used to visualizebiological relationships. These proceduresinclude a range of multivariate statistics suchas clustering techniques, multidimensionalscaling, and Mahalanobis’s generalized dis-tance. Craniometric analysis has been one ofthe transitional realms of osteological research.Pietrusewsky shows how this approach is stillappropriate for the investigation of widespreadmuseum collections, where destructive analysesare prohibited. Furthermore he demonstrates byusing examples from his own extensive researchin the Pacific, that multiple lines of evidence,including craniometric, dental, linguistic, andmolecular data are all necessary to contributeto our understanding of human populationhistory. Jackes tackles the problem of adultage determination and evaluates recent attemptsto circumvent some problems. She surveys andevaluates all of the different approachesto age-at-death estimation, including single

methods, such as metamorphosis of the pubicsymphysis and cranial suture closure, as wellas complex methods. She emphasizes the diffi-culties of dealing with the biases of referencesamples and the effects of skeletal preservationon efforts to produce age distributions forarchaeological samples. She takes the positionthat statistical investigation and manipulationcannot substitute for the necessity of havingaccurate biological age estimates. Finally,Milner, Wood, and Boldsen evaluate thecurrent status of paleodemography by focusingon some questions that have fueled pastdebates within the field. They address problemsof sampling, age and sex estimation, nonstatio-narity, heterogeneous risk, and selective mor-tality. Paleodemography draws from manyof the types of studies covered in previous chap-ters and attempts to tie together the success ofpopulations based on factors such as diet,disease experience, activity patterns, growthand development, and population interactions.Milner and colleagues provide a frank view ofthe potential and the limitations of achievingthe goal of being able to determine the level ofadaptation of past populations.

All of these chapters have the commontheme of determining information about pastpeoples from their skeletal and dentalremains. Adult age determination is an import-ant theme that appears in many chapters.Similarly, postmortem change, sampling, andthe relationship between cemetery samples andliving populations recurs throughout the book.Ethical considerations have had a majorimpact on all topics discussed. It is our hopethat this information will provide both breadthand depth for advanced studies in humanosteology and will serve as a guide to moreintensive study.

M. ANNE KATZENBERGSHELLEY R. SAUNDERS

PREFACE TO THE FIRST EDITION xix

ACKNOWLEDGMENTS

The editors wish to thank the contributors tothis volume for their efforts to present theirareas of research expertise to students andprofessionals. We are grateful to contributorswho have revised their work from the previousedition as well as those to who have preparednew contributions for this edition. It has

been a pleasure to work with everyone. Wealso thank our editors at Wiley, ThomMoore and Karen Chambers and our editorialassistant Ian Collins. Kristine Hepple andCharles FitzGerald provided excellenteditorial assistance during the final phases ofmanuscript preparation.

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CONTRIBUTORS

SABRINA C. AGARWAL, received her M.Sc.and Ph.D. (2001) from the University ofToronto, working in the Department ofAnthropology and the Samuel LunenfeldResearch Institute of Mount Sinai Hospital,Toronto. She completed a two-year SocialSciences and Humanities Research Councilof Canada Postdoctoral Fellowship in theDepartment of Anthropology, McMasterUniversity. She is currently assistant pro-fessor of anthropology and faculty associateat the Archaeological Research Facility,University of California, Berkeley. Herresearch interests focus broadly on thestudy of skeletal health and disease, withparticular focus on age, sex, and gender-related changes in bone quantity andquality. She is particularly interested in theapplication of biocultural and evolutionaryapproaches to the study of bone maintenanceand fragility. Her work has examined pat-terns of cortical bone microstructure,trabecular architecture, mineral density, andbiomechanical properties in several archaeo-logical populations in the OldWorld, includ-ing Medieval and Post-Medieval Britain,and Imperial Rome, as well as a pioneerpopulation from nineteenth-century UpperCanada. She is currently examining the

effects of growth and development and life-style on bone maintenance and fragilityover the lifecycle in an Anatolian Neolithicarchaeological population from Çatalhöyük,Turkey. She is also examining the long-term effect of parity and lactation on thematernal skeleton with a large-scale projecton bone maintenance in the nonhumanprimate (monkey) model. Her publicationsinclude a co-edited volume, Bone Loss andOsteoporosis: An Anthropological Perspe-ctive (Kluwer Plenum Academic Press)with S.D. Stout, published in 2003, and“Medieval Trabecular Bone Architecture:The Influence of Age, Sex and Lifestyle”American Journal of Physical Anthropology,2004, with M. Dimitriu, and M.D. Grynpas.

JESPER L. BOLDSEN, received his Ph.D. inbiology from the Department of TheoreticalStatistics, Aarhus University (Denmark) in1983. He is an associate professor with theInstitute of Forensic Medicine, Universityof Southern Denmark, Odense. He is alsoadjunct professor of anthropology at theUniversity of Utah and associate editor ofthe Journal of Biosocial Science andJournal of Applied Oral Science. Researchinterests revolve around human population

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biology, epidemiology, demography, andevolution. In recent years, he has concen-trated much of his research effort on analyz-ing the structure of the medieval populationof Denmark. This work has been based onstudies of extensively excavated medievalcemeteries. He has published widely inbiological and anthropological journals.Several recent publications include “Earlychildhood stress and adult age mortality—astudyof dental enamel hypoplasia in themed-ieval Danish village of Tirup,” AmericanJournal of Physical Anthropology, 2007;“Outside St. Jørgen: Leprosy in themedievalDanish cityofOdense,”American Journal ofPhysical Anthropology, 2006; “Leprosy andmortality in the Medieval Danish villageof Tirup,” American Journal of PhysicalAnthropology, 2005; and “Testing condi-tional independence in diagnostic palaeoepi-demiology,” American Journal of PhysicalAnthropology, 2005.

STEVEN K. BOYD, received his Ph.D. from theUniversity of Calgary in 2001. He thencompleted a postdoctoral fellowship at theSwiss Federal Institute of Technology,Switzerland, before joining the faculty atthe University of Calgary with a jointappointment in the Department of Mech-anical and Manufacturing Engineering(Schulich School of Engineering) and theFaculty of Kinesiology. Utilizing high-resolution computed tomography (micro-CT), his research has focused on the areaof orthopaedic biomechanics, focusing onadaptive changes to tissues that occur aftera joint injury or disease, with particular inter-est in bone. Recent publications include“Monitoring individual morphologicalchanges over time in ovariectomized ratsby in vivo micro-computed tomography,”Bone, 2006, with P. Davison, R. Müller,and J.A Gasser, and “Establishment ofan architecture-specific experimental vali-dation approach for finite element modelingof bone by rapid prototyping and high resol-ution computed tomography,” Medical

Engineering and Physics, 2006, with R. Suand G.M. Campbell.

JAMES BURTON, received his Ph.D. fromthe University of Arizona in 1986. He is thesenior scientist and associate director of theLaboratory for Archaeological Chemistryat the University of Wisconsin-Madison(Department of Anthropology). His researchinterests include the development of newarchaeometric methods, particularly the useof chemical and isotopic methods for prove-nience studies. Current projects includeexploration of alkaline-earth elements andvarious isotopic systems in the study ofhumanmobility and the development of non-destructivemethods to characterize historicalmaterials; studies of the effect of marineresources on bone levels of barium and stron-tium; the origins of the Gila Polychrome tra-dition; and the eastern, selvatic origin ofunusual pottery found at Late Formativesites in the Ecuadorian Andes. Recentlyhe co-authored with T.D. Price and V.Tieseler “Early African diaspora in colonialCampeche, Mexico: Strontium isotopicevidence” American Journal of PhysicalAnthropology, 2006, and “Interpreting thetrace-element components of bone—acurrent perspective from the Laboratoryfor Archaeological Chemistry” with T.D.Price in Deciphering Ancient Bones: TheResearch Potential of BioarchaeologicalCollections, edited by G. Grupe andJ. Peters (2003).

CHARLES M. FITZGERALD, received hisPh.D. in biological anthropology fromthe University of Cambridge in 1996. In2000, after completing a postdoctoral fellow-ship at McMaster University, he worked withSimon Hillson at the Institute of Archaeologyin University College London on a three-yearproject funded by the Natural EnvironmentResearch Council of the UK. This study’sobjective was to clarify the nature of thedental reduction that is observed in Middleand Upper Palaeolithic, Mesolithic, andEarly Neolithic hominids and to test the

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major hypotheses that account for it. Thestudy included the use of an approach toassess tooth size that overcame the problemof tooth wear, which was severe in theseearly modern humans. He returned toCanada where he was appointed the researchcoordinator for the Anthropology HardTissue and Light Microscopy Laboratory atMcMaster University. The focus of much ofhis research has been on the validation andapplication of odontochronological tech-niques, but in addition to growth and develop-ment, his interests embrace several other areasof skeleto-dental biology and palaeoanthro-pology. Recent publications include twopapers in the American Journal of PhysicalAnthropology: “Health of infants in anImperial Roman skeletal sample: perspectivefrom dental microstructure,” 2006, with S.R.Saunders, L. Bondioli, and R. Macchiarelli,and “A Test of histological methods of deter-mining the chronologyof accentuated striae indeciduous teeth,” 2005, with S.R. Saunders.

BENEDIKT HALLGRÍMSSON, received his Ph.D.from the University of Chicago. He is anassociate professor in the Departmentof Cell Biology and Anatomy, Faculty ofMedicine, University of Calgary, and anadjunct professor in the Departmentof Archaeology. He serves as the associatedean, Undergraduate Science Educationand is a member of the Alberta Bone andJoint Institute and an associate member ofthe Institute of Maternal and Child Health.Research interests include the developmental-genetic basis for phenotypic variationand the developmental-genetic basis forvariation in canalization, morphologicalintegration, and developmental stability.Dr. Hallgrimsson has published extensivelyin medical, biological, and anthropologicaljournals, including a co-edited volume,Variation: A Central Concept in Biologywith B.K. Hall (Academic Press, 2005).Recent articles include: “The brachymorphmouse and the developmental-geneticbasis for canalization and morphological

integration,” Evolution & Development,2006, with J.J.Y. Brown, A.F. Ford-Hutchinson, H.D. Sheets, M.L Zelditch,and F.R. Jirik, and “Canalization and devel-opmental stability in the Brachyrrhinemouse,” Journal of Anatomy, 2006, withK.E. Willmore, M.L. Zelditch, N. Young,A. Ah-Seng, and S. Lozanoff.

SIMON HILLSON, received his Ph.D. in Londonin 1979. He is a professor of bioarchaeologyat the Institute of Archaeology, UniversityCollege London. Dr. Hillson’s researchinterests focus on the biology of pasthuman populations, ranging from themost recent post-Medieval populations ofLondon to Predynastic Egyptians, andremains from Upper Palaeolithic contexts.In particular, he works with dental remains,studying their morphology, microstructure,development, pathology, and evolutionin Neanderthals and modern humans.As well as numerous journal articles,he has published three books: DentalAnthropology (Cambridge UniversityPress, 1996), Mammal Bones and Teeth(Institute of Archaeology, UCL, 1992) andTeeth (Cambridge University Press, 2005,second edition).

M. ANNE KATZENBERG, received her Ph.D. inanthropology from the University ofToronto in 1983. She holds a universityprofessorship at the University of Calgary(Department of Archaeology). Her researchinterests include diet and health in pastpeoples, and in particular, she explores thevarious applications of stable isotope analy-sis to reconstructing paleodiet, paleodemo-graphy, and ecology. She serves on theeditorial board of the Yearbook of PhysicalAnthropology and is the vice-president ofthe Paleopathology Association. In 2003,she was elected to the Royal Society ofCanada. She serves as a consultant inforensic anthropology for the MedicalExaminer of Alberta (southern division).Recent publications include “Identificationof historical human skeletal remains: a case

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study using skeletal and dental age, historyand DNA” International Journal ofOsteoarchaeology, 2005, with G. Oetelaar,J. Oetelaar, C. Fitzgerald, D. Yang, and S.R.Saunders, and “Skeletal Biology: GreatLakes Area,” Handbook of North AmericanIndians, Volume 3, 2006, edited byD.H. Ubelaker. She is co-editor,with Stanley Ambrose of Close to the Bone:Biogeochemical Approaches to PaleodietaryAnalyses in Archaeology (Plenum Press,2001).

ERIKA KRISTENSEN, received her B.Sc. in bio-logical engineering from the Universityof Guelph. She is completing herM.Sc. in the Department of Mechanicaland Manufacturing Engineering at theUniversity of Calgary and plans to start onher Ph.D. in late 2007. Her research interestsinclude applications of mechanical engine-ering principles to questions in developmen-tal biology, such as the relationship betweencraniofacial shape and mandibular strengthin growth-hormone-deficient mice. Sherecently submitted a paper titled “A novelhigh-throughput morphological method forphenotypic analysis,” with T.E. Parsons,B. Hallgrı́msson, and S.K. Boyd.

C. OWEN LOVEJOY, received his Ph.D. fromthe University of Massachusetts. He is auniversity professor in the Department ofAnthropology at Kent State University, aswell as serves in many other professionalcapacities, such as technical advisor inbiological anthropology, Cuyahoga CountyCoroner’s Office, Cleveland; adjunct pro-fessor of anatomy, Department of HumanAnatomy, NEOUCOM, Department ofOrthopaedic Surgery, School of Medicine,Case Western Reserve University). Hisresearch covers a broad spectrum of areasin human biology, from human paleo-anthropology to the genetics of develop-ment. Much of his research focuses onbipedal locomotion and its evolution. Manyof his publications involve the human post-cranium and that of our ancestors. Recent

articles include a series in Gait and Posture(2005–2006) on “The natural history ofhuman gait and posture”; and the article“Of muscle-bound crania and human brainevolution: The story behind the MYH16headlines,” Journal of Human Evolution,2006, with M.A. McCollum, C.C.Sherwood, C.J. Vinyard, and F. Schachat.

NANCY C. LOVELL, received her Ph.D. fromCornell University in 1987 and is a professorof anthropology at the University of Alberta.She specializes in bioarchaeology, paleo-pathology, and mortuary archaeology.Current research projects include thehistory of cemeteries in western Canadain order to assess the fit between archaeologi-cal interpretive models for prehistoriccemeteries with the documented evidencefor burial practices, and the analysis ofskeletal remains from ancient Egypt andMesopotamia to examine health and bio-logical relationships in complex societies.Recent publications include “Nonmetrictraits of the deciduous dentitions fromBronze Age Tell Leilan, Syria,” 2007, and“Nonmetric analysis of the permanentdentition of Bronze Age Tell Leilan,Syria,” 2006, both with S. Haddow andappearing in the International Journal ofDental Anthropology, and “A comparativeexperiment in the consolidation of crematedbone,” International Journal of Osteoar-chaeology, 2004, with D. Rossi andS. De Gruchy.

RICHARD S. MEINDL, received his Ph.D. fromthe University of Massachusetts. He isprofessor and chair of the Departmentof Anthropology at Kent State University.He conducts research in archaeology andosteology, forensic sciences, primate andhuman evolution, and historic demographyand population structure. Most recently, hehas studied the paleodemography ofvarious archeological areas, including theShell Mound Region of western Kentucky,a Late Woodland population from northernOhio, and a Late Classic site in lowland

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Yucatan. These areas provide specialopportunities to examine the rise ofcomplex societies, pre-agricultural sedentaryliving, and human demographic evolution.Publications include the review “RecentAdvances in Method and Theory inPaleodemography,” Annual Review ofAnthropology, 1998; and the co-editedbook Archaic Transitions in Ohio andKentucky Prehistory, with O.H. Prufer andS.E. Pedde (Kent State University Press,2001).

ROBERT P. MENSFORTH, received his Ph.D.from Kent State University. He joined thefaculty in the Department of Anthropologyat Cleveland State University in 1991. Hisresearch interests include skeletal biology,hominid paleontology, forensic anthropol-ogy, paleodemography, paleoepidemiology,medical anthropology, and human grossanatomy. Having completed the first phaseof a forensic study on patterns and types ofadult craniofacial trauma that characterize avery large sample of early twentiethcentury individuals from the Hamann–Todd collection at the Cleveland Museumof Natural History, his current project is theanalyses of the relationships among variousforms of trauma and the age, sex, andancestry of affected individuals. Among hispublications are “Paleodemography of theCarlston-Annis (Bt-5) late Archaic skeletalpopulation,” American Journal of PhysicalAnthropology, 1990, and “Warfare andtrophy taking in the Archaic Period,”Archaic Transitions in Ohio and KentuckyPrehistory, 2001, edited by O.H. Prufer,S.E. Pedde, and R.S. Meindl.

GEORGE R. MILNER, received his Ph.D. inanthropology from Northwestern Universityin 1982. He is currently professor ofanthropology at The Pennsylvania StateUniversity. Before coming to Penn State, hewas a postdoctoral fellow in physical anthro-pology at the Smithsonian Institution anddirector of the Museum of Anthropology atthe University of Kentucky. His research has

focused on both archaeology and humanosteology, with an emphasis on the prehistoryof eastern North America. He has conductedfieldwork on mortuary and habitation sitesin several midwestern states, Egypt, andthe Pacific. Currently he is serving onthe Editorial Board of the Journal ofAnthropological Archaeology. His latestbook is The Moundbuilders: AncientPeoples of Eastern North America (Thamesand Hudson, London, 2004) and, in 2006,The Cahokia Chiefdom: The Archaeology ofa Mississippian Society, was reprinted bythe University Press of Florida. Recent osteo-logical publications include a co-authoredarticle “Osteological applications of high-resolution computed tomography:Aprehisto-ric arrow injury,” Journal of ArchaeologicalScience, 2006, and “Skeletal biology:Northeast,” Handbook of North AmericanIndians: Environment, Origins, andPopulation, with Jane Buikstra and editedby D.H. Ubelaker (Smithsonian InstitutionPress, Washington, DC, 2006).

TRISH E. PARSONS, received her BSc. inbiological anthropology from ThePennsylvania State University. She iscurrently a student in the BiologicalAnthropology Graduate Program at theUniversity of Calgary. Her research interestsinclude the developmental basis of faciallength and the study of basic mammaliandevelopmental processes in the skullthrough mouse models and primateevolution. She has been a student ofmorphometrics since her days as anundergraduate at Penn State. Her mostrecent publication is “Microstructure oftrabecular bone in a mouse model forDown syndrome,” Anatomical Record:Advances in Integrative Anatomy andEvolutionary Biology, 2007, with T.M.Ryan, R.H. Reeves, and J.T. Richtsmeier.

MICHAEL PIETRUSEWSKY, received his Ph.D. inanthropology from the University of Torontoin 1969. He is currently professor of anthro-pology, University of Hawaii, Honolulu. His

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research focuses on studies of human skeletalremains from Australia, the Pacific, EastAsia, and Southeast Asia, including theapplication of multivariate statisticalprocedures to cranial data for investigatinghistorical–biological relationships. Recentpublications include “Ban Chiang, Aprehistoric village site in NortheastThailand II: The human skeletal remains,”The University Museum, University ofPennsylvania, Memoir Series, 2002, withM.T. Douglas; “Multivariate comparisonsof female cranial series from the RyukyuIslands and Japan,” AnthropologicalScience, 2004; “Matrilocality during the pre-historic transition to agriculture inThailand,” Antiquity, 2005, with R.A.Bentley, M.T. Douglas, and T.C. Atkinson;and “A multivariate craniometric studyof the prehistoric and modern inhabitantsof Southeast Asia, East Asia, andsurrounding regions: a human kaleido-scope?” Bioarchaeology of Southeast Asia,edited by M.R. Oxenham and N. Tayles(Cambridge University Press, 2006). He isa member of the American Association ofPhysical Anthropologists, fellow of theAmerican Academy of Forensic Sciences,diplomate of the American Board ofForensic Anthropologists, and member ofthe Editorial Boards of AnthropologicalScience, Anatomical Science International,and the Journal of Island and CoastalArchaeology. He has been a visitingprofessor/scholar to Australia, Canada,France, Germany, Japan, New Zealand,People’s Republic of China, and Taiwan.

DORI L. RAINEY, received her MA from theUniversity of Western Ontario (workingwith the skeletal remains of the Huron ofSouthern Ontario under Dr. MichaelSpence) and is currently working on herPh.D. in the Department of Anthropologyat McMaster University. With a broadfocus on skeletal biology, including palaeo-pathology and forensic anthropology, herresearch attempts to determine social

organization in archaeological populationsthrough nonmetric traits of the infracranialskeleton. More specifically, the researchendeavors to address the variability observedin nonmetric traits and to integrate these datawith pathological conditions, occupationalmarkers, and the archaeological context.

ALEXANDER G. ROBLING, received his Ph.D.from the University of Missouri in 1998and completed a postdoctoral fellowshipin the Department of Anatomy andOrthopedic Surgery at Indiana University—Purdue University, Indianapolis (IUPUI).He is currently an Assistant Professor in theDepartment of Anatomy & Cell Biologyand Biomedical Engineering at the IUPUI.Specializing in the effects of mechanicalloading on bone at the organ, tissue, cell,and molecular levels, his current interestsare focused on the signal transductioncascades involved in bone cell mechanosen-sation, including the genetic regulationof mechanosensitivity. Recent publi-cations include the co-authored article,“Biomechanical and Molecular Regulationof Bone Remodeling” Annual Review ofBiomedical Engineering, 2006; and thechapter, “Histomorphology, geometry, andmechanical loading in past populations”with S.D. Stout in Bone Loss andOsteoporosis: an AnthropologicalPerspective, (edited by S.C. Agarwal andS.D. Stout, (Kluwer Academic/PlenumPublishers, Inc.).

JEROME C. ROSE, received his Ph.D. in anthro-pology/biological anthropology from theUniversity of Massachusetts, Amherst, in1973. He is currently chair and professor ofanthropology at the University of Arkansas,Fayetteville, and a member of theDepartment of Anthropology, Institute ofArchaeology and Anthropology, YarmoukUniversity, Irbid, Jordan. His early researchinterest was dental anthropology with aparticular focus on dental histology, includ-ing the study of enamel microdefectsand enamel hypoplasias. Methodological

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