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Developmental Juvenile Osteology Second Edition Craig Cunningham Louise Scheuer Centre for Anatomy and Human Identification Centre for Anatomy and Human Identification School of Science and Engineering School of Science and Engineering University of Dundee, UK University of Dundee, UK Sue Black Centre for Anatomy and Human Identification School of Science and Engineering University of Dundee, UK Contribution by Helen Liversidge Department of Oral Growth and Development Queen Mary School of Medicine and Dentistry London University, UK Illustrations by Angela Christie AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 125 London Wall, London EC2Y 5AS, United Kingdom 525 B Street, Suite 1800, San Diego, CA 92101-4495, United States 50 Hampshire Street, 5th Floor, Cambridge, MA 02139, United States The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, United Kingdom Copyright © 2016, 2000 Elsevier Ltd. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Details on how to seek permission, further information about the Publisher’s permissions policies and our arrangements with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions. This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein). Notices Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility. To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein. Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library ISBN: 978-0-12-382106-5 For information on all Academic Press publications visit our website at https://www.elsevier.com/ Publisher: Sara Tenney Acquisition Editor: Elizabeth Brown Editorial Project Manager: Joslyn Chaiprasert-Paguio Production Project Manager: Lisa Jones Designer: Greg Harris Typeset by TNQ Books and Journals First Ed ition Foreword: Th e Dev elop ment of Juvenile Osteology The importance of this volume will be immediately live in the shadow of the perceived cutting-edge impor- obvious to anyone who has been confronted with fetal tance of these newer biochemical approaches to the or juvenile human osteological material in an archaeo- understanding and interpretation of skeletal material. logical, palaeontological, forensic or physical anthro- There is no doubt that these approaches are important, pological context. There is simply no currently available have enriched the disciplines that deal with the inter- reference work that deals with the fetal and juvenile pretation of skeletal material and have great potential human skeleton in sufficient detail to be practically to continue to do so. But there is still much to learn useful. Louise Scheuer and Sue Black have recognized and understand through the study of whole organisms. a major gap in the field and have responded with a Macro-anatomy gives much more than context and volume that is sure to become a classic wherever there background for these newer biochemical techniques. It is an interest in the identification and interpretation is here that this contribution will provide an invaluable of the human fetal and juvenile skeleton. and unparalleled resource. It is perhaps easy, or relatively so, to recognize a need Archaeologists, forensic pathologists and anthropolo- for a major reference work. It is much more difficult to gists will simply not be able to do without it in the con- fill that need. Both Scheuer and Black are highly expe- text of recognizing and identifying fetal and juvenile rienced anatomists with many years of classroom and material. It will also be invaluable to anyone interested research experience. There is no doubt about their qual- in human growth and development. For example, in my ifications to carry out this task. But even so, I doubt if at own field of human palaeontology, there is a growing the beginning they realized the enormity of the project, realization that there have been major changes in the they had set themselves or the length of time that would tempo and mode of ontogeny in human evolution and be needed to bring it to fruition. One major obstacle that a good understanding of these ontogenetic patterns was their conviction that the book must be based on will provide significant insight into our own evolution. skeletal material of known age to avoid the circularity To date, this work has focused primarily on the denti- of discussing age-specific skeletal development on the tion, but this volume will provide the necessary compar- basis of material that itself was aged using skeletal size ative context to facilitate the interpretation of skeletal or morphology. This proved to be difficult because such growth and development in pre-human fetal and juve- skeletal material is so rare and required considerable nile material. detective work to bring together. A second obstacle was Scheuer and Black have put considerable thought the wealth of previously published material scattered in into the organization of this work, making it not only many disparate references relevant to many different informative, but also accessible and practically usable. disciplines and published in many different languages. The meticulous descriptions of each individual bone The bibliography is large, spans 300 years and the infor- are clearly written and logically presented. The ‘practi- mation presented therein has been meticulously sorted cal notes’ for identification of all bones are invaluable and summarized. This in itself is a highly valuable con- to the field worker, as are the clear and beautifully exe- tribution to the field. The absolute insistence on doc- cuted illustrations by Angela Christie. There is no doubt umentation and accuracy of both the skeletal and the that this reference will outlive the current generation contextual information in the book will ensure that it of researchers. The authors and artist should be con- becomes a classic in the field. gratulated on providing a resource that will facilitate In recent times, descriptive anatomy has taken a defi- the research of so many current and future scientists nite back seat to the various biochemical approaches to whose work touches on the analysis of human fetal and skeletal analysis. Among others, these new approaches juvenile skeletal material. On behalf of all of us, I offer include DNA analyses that have the potential to uncover them a sincere and well-deserved thank you. the genetic basis of skeletal growth, the sex and possible familial and/or ethnic affiliation of skeletal material Professor Leslie C. Aiello and the infectious diseases that the individual suffered President, Wenner-Gren Foundation for in life. There is also the possibility, through stable iso- Anthropological Research, Inc., New York; tope analysis, of determining the diet of the individual. Emeritus Professor, University College London; In many academic departments, topographic anatomists Honorary Fellow, University College London vii Seco nd Ed ition F orewo rd It is difficult to believe it is 16 years since the publica- and development. There is also tantalizing evidence that tion of the first edition of what is known in the trade as the histology of fossil bone may also provide a window ‘Scheuer and Black’. into growth and development. These advances, plus the First, ‘hats off’ to the editor who managed to persuade reality that hard tissue research is even more special- the powers that be to commission a second edition of ized now than it was in 2000, have been recognized in this important reference book. Academic publishers pre- the second edition of Developmental Juvenile Osteology by pared to look further than the next budget cycle are as inclusion of a separate chapter contributed by Helen rare as hen’s teeth, so Elsevier’s investment in the future Liversidge on the dentition. of Developmental Juvenile Osteology is a welcome reversion The original authors, now joined by a new senior to the traditional values of academic publishing. author, Craig Cunningham, have obviously thought In the Foreword to the first edition, Leslie Aiello set long and hard about how to improve an instant ‘clas- out the case for a book about the developing modern sic’, and their decisions about what to leave well alone human skeleton. There is clearly a forensic need to and what to change are well-judged. Plus two of the identify immature skeletal remains. But there is also a great strengths of the first edition, the illustrations and need for a book that reminds anatomists, and especially the copious bibliography, have been, respectively, aug- paleoanthropologists, that skeletal ontogeny has a lot mented and updated. to tell us about human evolution. Few juveniles seem to The first edition of Henry Gray’s human anatomy make it into the fossil record, but I suspect that some book was published in 1858. A second edition rapidly juvenile fossils go unrecognized because most research- followed in 1860, but this was to be the last by Gray for ers are programmed to look for, and to recognize, the he died a year later of smallpox. Thankfully, Louise remains of adults. In this respect ‘Scheuer and Black’ Scheuer and Sue Black are in rude health. By my cal- has helped educate our community. culation they are 142 years behind Henry Gray, but as One of the many ways modern humans differ from his initial vision lives on, now so does theirs. May it long our closest relatives, chimpanzees and bonobos is the continue to do so. rate at which we progress through the important devel- opmental way stations we call life history. The hard tissue Professor Bernard Wood fossil record is a potential window into the evolutionary University Professor of Human Origins, history of these life history differences, and one of the George Washington University, Washington DC, USA; most important advances in paleoanthropology in the Honorary Professor, University of past several decades has been the exploitation of dental Kent Canterbury, UK microstructure to calibrate pre- and post natal growth ix Preface In the 16 years since the publication of the first edi- the first edition, most have been sourced directly from tion of Developmental Juvenile Osteology, much has actual bone specimens housed in what has become advanced in the field of skeletal development, with a widely known as the Scheuer collection. resurgence of interest in the growing skeleton. This, As stated in the first edition, it has always been the combined with new published literature, has greatly intention that Developmental Juvenile Osteology would be enhanced our understanding of the human skeleton. primarily aimed at skeletal biologists, forensic anthro- This revised edition has attempted to bring together pologists, forensic pathologists, archaeologists and this new and diverse array of research literature to pro- palaeontologists but it is hoped that this revised edi- vide an updated account of skeletal development and tion may perhaps prove both interesting and useful to a to furnish the reader with an expanded bibliography. wider clinical and scientific audience. Indeed, a strength of the first edition was its extensive Finally, thanks must be conveyed to those who have bibliography and this edition adds more than 1000 new assisted throughout the production of this edition and references, documenting many of the significant contri- our gratitude remains with those previously listed in the butions to the field in the intervening period. first edition. However, thanks are extended to Helen The main core of the text persists by describing each Liversidge, an acknowledged authority on tooth devel- individual component of the human skeleton from its opment and ageing, who has compiled a new chapter embryological origin through to its final adult form. on the dentition. Lucina Hackman, Paul Felts and This systematic approach has been shown to assist the Catherine Carr are acknowledged for their contribu- processes of both identification and age estimation and tions to sections on age estimation in the living, bone acts as a core resource for the basic understanding of histology and embryology, respectively. Gratitude is normal human skeletal development. In addition, new extended to Joslyn Paguio, Elizabeth Brown and Lisa sections have been introduced where there have been Jones from Elsevier who have looked after the produc- significant advances in the field, including dental devel- tion of this revised edition and have provided their opment and age estimation from living individuals. support and encouragement throughout the revisions. Throughout the text, data tables have been updated Thanks are also given to Christopher Rynn who assisted with the most relevant and up-to-date information. Easy- in the design of the front cover and to Katie Tyldesley to-read compilation summaries of key skeletal events who completed initial literature searches in the early and associated literature have also been expanded. stages of this revised edition. Additionally, material that was presented in Developmen- tal Juvenile Osteology’s sibling texts: The Juvenile Skeleton Craig Cunningham and Juvenile Osteology: A Laboratory and Field Manual Louise Scheuer have been incorporated into this edition. New illustra- Sue Black tions and schematics have been introduced and as with xi C h a p t e r 1 Introduction: A Guide to the Text It must surely be clear that if we wish to safeguard the interests The first edition of this text was published in 2000 of our science (physical anthropology), and of those innocents and in the intervening period there has been a resur- who identify themselves with it, and who by so doing volun- gence of interest in this fascinating field, but the tarily condemn themselves to a precarious, albeit interesting main purpose of this book remains the same. Its aim life brachiating as it were from one lower income bracket to is to describe each individual bone of the skeleton, or another, then it is our duty to see to it that they are properly indeed different components of a bone, and follow equipped for the work which they wish to do and which so development from embryological origin through to urgently requires to be done. the final adult form. This systematic approach has been (Montagu (1941)) shown to assist the processes of both identification and T age estimation of the juvenile skeleton and to aid as a he correct identification of the skeletal components core source for basic understanding of normal human of the juvenile skeleton is critical to the analysis of skel- skeletal development. The passage of time has ensured etal remains, regardless of whether they are of archaeo- that the already full reference section is enhanced fur- logical or forensic origin. Without such information it is ther and although the main core of the text persists, virtually impossible to establish the number of individu- new sections have been added where it was felt that the als represented, let alone ascertain their identity and expertise of others would enhance the original concept. make meaningful inference on demographic or health- Chapter 2–4 form a general introduction to related issues. Indeed, a lack of familiarity with immature the juvenile skeleton. Chapter 2 considers many of remains has led, on more than one occasion, to their the fundamental issues concerning juvenile skeletal classification as ‘non-human’ or ‘of uncertain origin’. remains including the origin of such material, the vari- Once the remains have been confirmed as human, the ous techniques by which it has been studied, the vari- next step is usually an attempt to establish the four prin- ability of child growth, the dilemma of biological versus cipal parameters of biological identity (sex, age at death, chronological age and skeletal versus dental age. Age stature and ancestry). However, with juvenile skeletal estimation in the living has become a major issue for remains it is often only the estimation of the age at death border control, migration and human trafficking, and that can be established with any degree of reliability. Sex our thanks are extended to Dr. Lucina Hackman who estimation from juvenile remains is tentative at best, and has introduced this subject here to demonstrate that stature is so closely linked to the age of the individual that age determination from the skeleton is not restricted it is often used to predict it. Ancestry is difficult to estab- to the domain of the deceased. Chapter 3 examines lish in the adult, so in the child it is highly speculative at the more specific cellular and vascular nature of bone best, especially when only skeletal remains are presented. growth and development. It discusses the ontogenetic The value of understanding juvenile skeletal remains development of bone from its mesenchymal origins, should not be overlooked in both the medical and the through a cartilaginous or membranous template, to anatomical fields of study. A glance through the ref- its eventual transformation into bone. Bone growth erence section will demonstrate the critical nature of is considered, as is the influence of its vascularity. Dr. understanding normal juvenile development to facili- Paul Felts enhances the value of this section not only in tate diagnosis and treatment of aberrant conditions. relation to the basic cellular composition but also the Developmental Juvenile Osteology. http://dx.doi.org/10.1016/B978-0-12-382106-5.00001-3 Copyright © 2016 Elsevier Ltd. All rights reserved. 1 Developmental Juvenile Osteology relevance to bone modelling and remodelling as it per- previously in Chapter 4 and deals with the specific tains to not only normal growth but also to repair and embryological and early fetal development of that par- pathology. Chapter 4 has been reviewed and updated by ticular bone. This section charts its development from Dr. Catherine Carr and provides a very brief outline the blastemal condition up to the stage prior to the com- of the early embryological development of the human mencement of ossification. It also includes reference to body as a whole, and sets the scene for the more spe- various congenital conditions and anomalies that may cific developmental aspects of the skeleton that are dis- arise during this period and which could subsequently cussed in subsequent chapters. alter the final adult morphology of the bone. Chapters 5–12 respectively form the core of the text The section on ossification describes the develop- and describe the morphological development of the ment of the bone from the time of appearance of the immature skeleton in a way that permits the ready iden- first centre(s) of ossification up to the stage of final tification of each skeletal element and thus allows an fusion of the epiphyses. In most chapters, this section evaluation of the age at death of the individual. The is in three parts: primary centres, secondary centres chapters are arranged in a topographical order, com- and pattern of epiphyseal fusion. It is in this section mencing with the axial skeleton and continuing with of the book that the illustrations are most important the upper and then the lower limb girdles and their as they not only highlight the earliest stage at which a associated appendages. Each chapter is essentially sepa- particular element can be identified with certainty, but rated into four sections–the adult bone, early develop- also describe the morphological changes that occur ment, ossification and practical notes. Chapter 6 takes in that bone throughout its development. However, a different format as it considers the dentition, and Dr. it has not always been possible to illustrate a specific Helen Liversidge, has greatly enhanced this section stage of development due to the limited availability of from the original text and included the latest data on material. The illustrations in this section are half-tone ageing from the dentition. drawings of bones, many of which are of known age Each of the main chapters begins with a description and sex and again only the right side of the body is of the adult bone(s) but this is far from an exhaustive depicted. It should be mentioned that all of the draw- consideration of the subject as there are many excellent ings from Mrs. Angela Christie are derived from actual texts written specifically to fulfil this purpose. However, bone specimens which are now housed in the Scheuer it was deemed necessary to include this section primar- collection at the University of Dundee. As an active ily to ensure consistency of terminology used in the repository for juvenile skeletal remains, it is an invalu- subsequent sections on the development and ossifica- able source but it cannot in any way be considered as a tion of the bone. Where possible, the accepted standard ‘population’ as the origins of the material are diverse– anatomical planes and terminology have been used anatomical specimens, archaeological material and throughout, although more commonly used names and forensic casework. others that reflect a historical origin have sometimes The final section within each chapter is headed been included. Several anomalies of the adult skeleton Practical Notes. This represents a summary or mor- have been addressed as the understanding of variation phological timetable of critical osteological events from is an important concept that is diminishing as teaching the commencement of ossification to final epiphyseal of anatomy in particular moves away from more tradi- fusion (or the attainment of final adult form). The tional methods towards computer models and plastic practical notes include guidelines on the sideing of skeletal teaching aids. While it is appreciated that these remains and how to orientate them to achieve correct minor skeletal variants may be of limited clinical value, identification of the skeletal element. In addition, there they can occasionally prove extremely important in the is a small section that offers suggestions on which other identification of the deceased. In anthropological terms bones that have a similar morphology may cause some of course, many of these anomalies are referred to as confusion and thus result in misidentification. non-metric traits that may be considered indicative of Finally, some tables of metric information are potential genetic influences (Berry, 1975; Finnegan, included that may prove useful in the estimation of age 1978). A variety of relevant clinical conditions has also at death. This tends to include primarily observations been introduced in this section where they have some on individuals of documented age to remove the inherent bearing on the future development of the bone. Com- errors of the circular argument that ensues when age is ment has often been made with regards to the value subsequently predicted on the basis of the accuracy of of that particular element in the assessment of some another method (see Chapter 2). Naturally, this reduces parameters of biological identity (sex, ancestry and the number of studies that could be included but it may stature). While this is not the primary aim of this text, serve to highlight where further research could be pur- it serves only to direct the reader to other sources of sued. The additional sources of information collated reference. for The Juvenile Skeleton (Scheuer and Black, 2004) and The illustrations of the adult bones are represented for Juvenile Osteology (Schaefer et al., 2009) have been by stippled line drawings with muscle attachments indi- included in this second edition. cated. The illustrations throughout the book always By far the most comprehensive account of fetal bone depict the right-hand side of the body. remains is that published by Fazekas and Kósa (English The early development of each bone is described translation of 1978) referring to a group of 136 fetuses directly after the discussion of the adult morphology. ranging from 12 to 40 weeks gestation. However, the Each description follows on from the stage outlined sample was essentially of undocumented age at death 2 Chapter 1 Introduction: A Guide to the Text and age was assigned on the basis of its well-documented of the most basic descriptions of bones are historical relationship with body length (Streeter, 1920; Scammon and are where we tend to find the greatest attention to and Calkins, 1929; Schultz, 1929a). There is, however, no detail, so we make no apology that the references span other detailed text on fetal osteology and given the fact over 300 years. Wise (1995) accused many authors of that all fetal material must, by necessity, be of uncertain ignoring the contributors of the past, stating that ‘we age (see Chapter 2) its inclusion remains justified. may have stood on the shoulders of giants but we did As each bone of the skeleton is considered from its not cite them’. He attributes this to authors becoming earliest formation to its adult morphology, it is obvious victims of technology, relying on the use of informa- that each would display its own idiosyncrasies and resist tion retrieval systems that tend not to extend to more being forced into a standard chapter format. As a result, than 25 years ago–but that does not mean that they while an attempt has been made to adhere to an orga- should be consigned to obscurity because of modern nized structure, each chapter is, by necessity, slightly laziness that keeps us in front of our screens and does different in terms of its layout. For example, in Chap- not see us resort to the original source. Many of the ter 5 (Skull) there is a general introduction to the early older texts may also express views and descriptions development of the skull as a whole to prevent need- that would not now be considered ethically accept- less repetition of material that is common to a structure able. Titles of these papers have obviously been given composed of so many conjoined elements. Also, there is as they stand and, where appropriate, the text has no section on secondary centres, as these do not occur been quoted verbatim in the hope that accusations of in the skull. Similarly in Chapter 7, as the vertebral col- political incorrectness may not be directed at this text. umn is an axial structure, there is obviously no section O’Rahilly (1996) raised the criticism that rather than on side identification and instead this is replaced by seek out the original reference, many authors substi- identification of position within a series. tute reviews of the subject or even cite student text- In addition to the principal elements of the skeleton, books where, in all fairness, original research is rarely other structures such as the costal cartilages have been published. In addition, he further accused authors of included. Being composed of hyaline cartilage, these repeating information from one text to another with- structures maintain the potential to ossify and may do out due recourse to the original work, which can of so at an age when the remainder of the skeleton is still course lead to the perpetuation of errors. Where pos- in its late developmental phase. For this reason it is sible this text has remained true to its original ethos important that the structures can be identified, as they and attempted to avoid these pitfalls by extensive lit- may be encountered in the excavation or retrieval of erature searches with recourse to the original texts immature remains. While such ossifications have always but it is inevitable that vital references, perhaps in tended to be considered entirely within the domain of another language, may have been omitted and errors the elderly, the inaccuracy of this assumption is high- may indeed have been unwittingly perpetuated. It is lighted and awareness of their existence can lead to an hoped that readers may identify these and make them increase in successful retrieval rates. known to the authors of this text so that due rectifica- One of the overwhelming comments to arise from tion can be made. the first edition of this text was the value of the exten- A second edition gives the opportunity to address sive reference section which has been expanded in errors and omissions in the first text, and we are grateful this edition to ensure currency. The quantity of litera- to those who pointed out where they occurred and we ture differs for each bone and so by necessity some hope this new edition will have corrected those that were areas are more heavily referenced than others. Many most obvious and not introduced too many new ones. 3 C h a p t e r 2 Skeletal Development and Ageing Chapter Outline Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Source Material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Estimation of Skeletal Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Estimation of Dental Age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Age Estimation in the Living . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Skeletal Remains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Documentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Sampling and Representativeness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Sex Estimation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 The childhood shows the man It is equally important that forensic scientists can As morning shows the day also recognize juvenile components of the skeleton and (John Milton, Paradise Regained) establish a reliable age at death to assist in determin- B ing, or to confirm, the identity of an individual. Foren- iological identity is used for different purposes sic anthropology uses the biological identity to assist in depending on the aims of the investigator. Skeletal biolo- determining ‘personal’ identity and this demands accu- gists, physical anthropologists and palaeodemographers racy and reliability, as it is only when the deceased has use the information to construct demographic profiles of been given a name that an investigation can proceed. populations from both historic and prehistoric times and This chapter considers the basic concepts of growth then draw conclusions about lifestyle, death rates and life and its relationship to age and then attempts to distin- expectancies. In a skeletal assemblage that includes sub- guish the many different categories of database from adult specimens, the identification and age at death of which information on the developing skeleton is drawn. the juvenile component will be particularly relevant as It then summarizes the estimation of age from skeletal it is deemed a reflection of the overall health and well- and dental material and ageing from the living. Finally, being of that population. aspects of documentation, sampling, representativeness Developmental Juvenile Osteology. http://dx.doi.org/10.1016/B978-0-12-382106-5.00002-5 Copyright © 2016 Elsevier Ltd. All rights reserved. 5 Developmental Juvenile Osteology and sexing that are particularly associated with the juve- allows more growth beforehand and therefore has its nile component of a skeletal assemblage are discussed. greatest influence at a different critical phase of growth. It results in a greater adult size, predominantly because muscle mass increases rapidly during this period, which GROWTH affects overall skeletal robusticity (Tanner, 1978). Some studies show that, as in childhood, bone mineral density Growth is a term that is used to describe progressive and the rate of accumulation of peak bone mass vary changes in size and morphology during the development between the sexes during puberty (Gordon et al., 1991). of an individual. In general, it is correlated positively with However, Baxter-Jones et al. (2003) have questioned the age and so estimation of age at death utilizes the many significance of this. incremental changes that occur during development. Genetic differences are the basis for differences Growth consists of two components, increase in size and between population groups and it is self-evident that increase in maturity, and while these two elements are the adults of different genetic ancestry have overall size closely related, their relationship is not linear. As a result, differences; witness a group of Japanese and a group individuals reach developmental milestones, or bio- of Dutch tourists. A comprehensive survey of variation logical ages, along the maturity continuum at different in the growth of children worldwide can be found in chronological ages. For example, two boys, both aged Eveleth and Tanner (1990). The difference that the 8 years, may differ considerably in height or similarly, environment may make on this intrinsic genetic factor two girls, aged 13, may both be at sexually and skeletally is complex, and one interesting approach to its eluci- different stages of maturity. Generally speaking, growth dation has been longitudinal growth studies on mono- in size is a regular process, although there are distinct zygotic and dizygotic twins and their siblings (Tanner, increases in rate, between 6 and 8 years in mid-childhood 1962). By far the most important environmental fac- and at the adolescent growth spurt. However, the only tors are those that can be grouped together under the consistent characteristic of growth is its variability. umbrella of socio-economic influences and they may There are variations in growth rate between differ- be subdivided into the effects of nutrition, disease and ent tissues and organs of the body in all individuals. social status itself. Although all of these factors dominate The brain and head attain their adult size early in most strongly in infancy and childhood, their extremes childhood, while the lymphoid system reaches its peak can affect growth and development even before birth. in late childhood. The reproductive system displays The starvation conditions in both Russia and the Neth- yet another rate and develops later in the adolescent erlands during World War II caused a significant decline period (Tanner, 1978). Growth also varies between the in both birth weight and vitality of infants (Antonov, sexes, between individuals of the same population and 1947; Smith, 1947). Similarly, research following women between populations themselves. The underlying basis who were pregnant during the 1998 Quebec ice storms of this variation is primarily genetically determined has shown that prenatal maternal stress has long-term but the influence of environmental factors is critical in consequences on the child (Cao-Lei et al., 2014). The controlling the expression of the growth process. This effects on optimum size and weight at birth and poor interplay between genetic and environmental influ- growth in the early years have been shown to affect not ences is the basis of the ‘nature versus nurture’ argu- only final adult size, but also susceptibility to disease ment. In spite of much research, the causal picture still (Frisancho et al., 1970; Clark et al., 1986; Barker et al., remains unclear. On the one hand, it is almost impos- 1993). Maternal undernutrition appears to be one of sible to study the effects of a single factor alone and the links in the causal chain between socio-economic on the other, the effects of a factor on an individual factors and fetal growth (Lechtig et al., 1975). Nutrition may vary, depending at which stage of development it and disease have long been accepted as factors in raised acts. Thus the causes responsible for differences in any childhood morbidity and mortality rates in countries particular person are complex and difficult to isolate. with low socio-economic levels. Even in countries with Genetic inheritance is the background for differences a general high standard of living, minority groups with of size and maturity between the sexes, which, although a lower than average income, raise children who show small, can be discerned, even before birth (Choi and delayed postnatal ossification rates and tooth emer- Trotter, 1970; Pedersen, 1982). These manifest in the tim- gence times (Garn et al., 1973a,b). ing of ossification of bones and mineralization of teeth Minor factors such as season and climate are also (Garn et al., 1966a; Mayhall, 1992). Postnatally, skeletal thought to affect rates of growth and maturity. It is maturation is more advanced in girls than boys (Pyle and known that growth in height and weight differs accord- Hoerr, 1955; Brodeur et al., 1981) but bone mineral den- ing to the time of year but it is debatable as to what sity is significantly less in girls than boys, the latter having extent climate affects these processes. Studies com- a higher mineral density and larger long bones (Maresh, paring ethnic minorities in their adopted countries to 1970; Specker et al., 1987; Miller et al., 1991). As puberty populations still living in their original homeland have approaches, sexual dimorphism increases by differential often given contradictory results. Again, it is difficult to hormone secretion and this is reflected in the adoles- isolate climate from the many changes that have acted cent growth spurt. Although the timing varies between on the immigrant community. individuals of the same sex (early and late maturers), Evidence has accumulated that over the past 150 years girls are, in general, about 2 years in advance of boys in there has been an increase in height and weight of adults maturity at the same age. The later growth spurt in boys and a decrease in the age at which adult size is achieved 6

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